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1.
Using a free-air CO2 enrichment (FACE) experiment, poplar trees (Populus × euramericana clone I214) were exposed to either ambient or elevated [CO2] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm−2) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (g s, μmol m−2 s−1), photosynthetic CO2 fixation (A, mmol m−2 s−1), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO2 (Ci:Ca). Elevated [CO2] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO2] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO2], g s was reduced, A/E was stimulated, and Ci:Ca was reduced relative to ambient [CO2]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO2]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO2], although the mechanism for this remains obscure.  相似文献   

2.
We studied the effects of repeated defoliation and drought, which are considered to be most effective in triggering oak decline, on the Central-European oak species Quercus robur L. and Quercus petraea [Matt.] Liebl. Three-year-old saplings were subjected to artificial defoliation in two consecutive years and to drought stress, applied singly or in combination. Differences in the morphological, anatomical and physiological responses to these stress factors among treatments and between species were tested in a 3-year experiment. Drought stress lowered the pre-dawn and the afternoon leaf-water potentials in both species, irrespective of additional defoliation. The relative water content of the leaves was unaffected by drought stress; but, in Q. robur, it decreased in trees subjected to defoliation and to the combination of drought stress and defoliation. When repeated defoliation and drought stress were applied within the same growing season, the leaf-to-air difference in the partial pressure of water vapour (w) explained most of the variation in the daily integral of the stomatal conductance to water vapour (g s day sum) in Q. petraea; however, in Q. robur, the hydraulic conductance of the twigs was the most powerful predictor variable. In the same year, the actual hydraulic conductance was correlated with g s day sum in Q. robur, but not in Q. petraea. The studied parameters of gas exchange and biomass production revealed that Q. robur recovered more rapidly from stress than did Q. petraea. We hypothesize that this is due to Q. roburs lower responsiveness to the environmental variable w and its higher responsiveness to internal hydraulic conductance compared to Q. petraea. This revised version was published online in May 2005 with corrections to Table 4.  相似文献   

3.
Pedunculate (Quercus robur L.) and sessile (Q. petraea [Matt.] Liebl.) oaks are the most common oak species in Western Europe. They are known to display different ecological requirements, particularly relative to root hypoxia induced by flooding: In a glasshouse study of seedlings, we quantified the effects of flooding on starch mobilization from cotyledons and starch partitioning. Growth and distribution of lateral roots were also measured. The above-ground growth of Q. robur was less affected by flooding than that of Q. petraea which failed to develop a second flush. Root growth was also severely inhibited, particularly in Q. petraea. In Q. robur, lateral root initiation as well as elongation was restricted to the soil surface layer. Flooding markedly reduced total growth and concentrations of in all components except stems. Starch mobilization from cotyledons was delayed by flooding, especially in Q. robur seedlings. Under flooding, the decrease of cotyledons dry mass and starch content in Q. robur was lower than in Q. petraea, whereas Q. robur displayed larger growth than Q. petraea. The features of carbohydrate management may be crucial in the observed differences in flooding tolerance of these species.  相似文献   

4.
Background and Aims The inverse relationship between atmospheric CO2 partial pressure (pCO2) and stomatal frequency in many species of plants has been widely used to estimate palaeoatmospheric CO2 (palaeo-CO2) levels; however, the results obtained have been quite variable. This study attempts to find a potential new proxy for palaeo-CO2 levels by analysing stomatal frequency in Quercus guyavifolia (Q. guajavifolia, Fagaceae), an extant dominant species of sclerophyllous forests in the Himalayas with abundant fossil relatives.Methods Stomatal frequency was analysed for extant samples of Q. guyavifolia collected from17 field sites at altitudes ranging between 2493 and 4497 m. Herbarium specimens collected between 1926 and 2011 were also examined. Correlations of pCO2–stomatal frequency were determined using samples from both sources, and these were then applied to Q. preguyavaefolia fossils in order to estimate palaeo-CO2 concentrations for two late-Pliocene floras in south-western China.Key Results In contrast to the negative correlations detected for most other species that have been studied, a positive correlation between pCO2 and stomatal frequency was determined in Q. guyavifolia sampled from both extant field collections and historical herbarium specimens. Palaeo-CO2 concentrations were estimated to be approx. 180–240 ppm in the late Pliocene, which is consistent with most other previous estimates.Conclusions A new positive relationship between pCO2 and stomatal frequency in Q. guyavifolia is presented, which can be applied to the fossils closely related to this species that are widely distributed in the late-Cenozoic strata in order to estimate palaeo-CO2 concentrations. The results show that it is valid to use a positive relationship to estimate palaeo-CO2 concentrations, and the study adds to the variety of stomatal density/index relationships that available for estimating pCO2. The physiological mechanisms underlying this positive response are unclear, however, and require further research.  相似文献   

5.
Pedunculate oak (Quercus robur L.) was germinated and grown at ambient CO2 level and 650 ppmv CO2 in the presence and absence of the ectomycorrhizal fungus Laccaria laccata for a total of 6 month under nutrient non-limiting conditions. Mycorrhization and elevated atmospheric CO2 each supported the growth of the trees. Stem height, stem diameter, and dry matter accumulation of pedunculate oak were increased by mycorrhization. Elevated atmospheric CO2 enhanced stem height, stem diameter, fresh weight and dry weight, as well as lateral root formation of the trees. In combination, mycorrhization and elevated atmospheric CO2 had a more than additive, positive effect on tree height and biomass accumulation, and further improved lateral root formation of the trees. From these findings it is suggested that the efficiency of the roots in supporting the growth of the shoot is increased in mycorrhized oak trees at elevated atmospheric CO2.Abbreviations DW dry weight - FW fresh weight - RWC relative water content  相似文献   

6.
Photosynthetic acclimation of two co-occurring deciduous oaks (Quercus petraea and Quercus pyrenaica) to a natural light gradient was studied during one growing season. In the spring of 2003, 90 seedlings per species were planted along a transect resulting from a dense Pinus sylvestris stand, an adjacent thinned area and a 10-m-wide firebreak (16.5–60.9% Global Site Factor (GSF)). In two dates of the following summer, we measured leaf gas exchange, carboxylation efficiency (CE), chlorophyll and nitrogen content, light–response curves of chlorophyll a fluorescence parameters, and leaf mass per area (LMA). Summer was mild, as evidenced by leaf predawn water potential (Ψpd), which reduced the interactive effect of water stress on the response of seedlings to light. Q. pyrenaica had higher LMA, CE, stomatal conductance (g s max) and photosynthesis per unit area than Q. petraea at all growth irradiances. , LMA, g s max and electron transport rate (ETR) all increased with light availability (GSF) in a similar fashion in both species. Light had also a clear effect on the organization of Photosystem II (PS II), as deduced by chlorophyll a fluorescence curves. Chlorophyll concentration (Chlm) decreased with increasing light availability in Q. pyrenaica but it did not in Q. petraea. Seedlings of Q. petraea acclimated to higher irradiances showed a greater non-photochemical quenching (NPQ) than those of Q. pyrenaica. This suggests a higher susceptibility to high light in Q. petraea, which would be consistent with a better adaptation to shade, inferred from the lower LMA or the lower rate of photosynthesis.  相似文献   

7.
The response of stomata in isolated epidermis to the concentration of CO2 in the gaseous phase was examined in a C3 species, the Argenteum mutant of Pisum sativum, and a crassulacean-acid-metabolism (CAM) species, Kalanchoë daigremontiana. Epidermis from leaves of both species was incubated on buffer solutions in the presence of air containing various volume fractions of CO2 (0 to 10000·10–6). In both species and in the light and in darkness, the effect of CO2 was to inhibit stomatal opening, the maximum inhibition of opening occurring in the range 0 to 360·10–6. The inhibition of opening per unit change in concentration was greatest between volume fractions of 0 and 240·10–6. There was little further closure above the volume fraction of 360·10–6, i.e. approximately ambient concentration of CO2. Thus, although leaves of CAM species may experience much higher internal concentrations of CO2 in the light than those of C3 plants, this does not affect the sensitivity of their stomata to CO2 concentration or the range over which they respond. Stomatal responses to CO2 were similar in both the light and the dark, indicating that effects of CO2 on stomata occur via mechanisms which are independent of light. The responses of stomata to CO2 in the gaseous phase took place without the treatments changing the pH of the buffered solutions. Thus it is unlikely that CO2 elicited stomatal movement by changing either the pH or the HCO 3 /CO 3 2- equilibria. It is suggested that the concentration of dissolved unhydrated CO2 may be the effector of stomatal movement and that its activity is related to its reactivity with amines.  相似文献   

8.
植物化石气孔参数分析是目前恢复古大气二氧化碳浓度较为精准的方法之一,银杏类和松柏类等是恢复古大气CO_2浓度常用的化石类群。本文利用新疆准噶尔盆地下侏罗统三工河组的松柏类掌鳞杉科Brachyphyllum(Hirmeriella?)sp.化石对早侏罗世大气CO_2浓度进行了重建,获得早侏罗世大气CO_2浓度为~1200ppm,丰富了早侏罗世大气CO_2浓度信息,进一步说明掌鳞杉科植物通过气孔比率法在重建侏罗纪大气CO_2浓度方面的可靠性。掌鳞杉科植物的旱生构造和较高的大气CO_2浓度表明早侏罗世Toarcian期大洋缺氧事件在陆地生态系统内可能产生了一定的响应。  相似文献   

9.
Stomatal frequency is often observed to vary inversely with atmospheric CO2 concentration (pCO2). The response is due to (1) individual phenotypic plasticity and (2) evolutionary change, depending on the time scale. Evolutionary responses occur more frequently than individual responses and individual responses are more pronounced under subambient pCO2 levels than under elevated pCO2 (CO2 ceiling). The evolutionary response appears therefore to be a valuable device for determining past pCO2. Since tree leaves often represent a conspicuous and rich resource of fossil material, they are increasingly important in this respect. Additionally, certain tree species are considered to represent living fossils and therefore valuable sources of ancient stomatal data. There are, however, numerous difficulties which have to be considered such as: (1) high variance of the data, especially for fossil material, (2) interspecific differences of the response, (3) the CO2 ceiling and (4) differences between short-term and long-term responses. Whereas the qualitative pCO2 signal of stomatal frequency appears to be reliable, quantitative pCO2 reconstruction has to be performed with caution. The results of a number of studies which used stomatal frequency as a pCO2 sensor demonstrate good agreement with the results obtained with other proxy data. Current techniques are based on transfer functions which calibrate the fossil data with extant material. It is suggested that a mechanistic approach including physical as well as physiological processes could improve pCO2 reconstruction. Furthermore, the topic of the influence of pCO2 on stomatal frequency is significant not only for reconstructing past pCO2 but also with respect to the climate-biosphere interrelationship.  相似文献   

10.

Background and Aims

The inverse relationship between stomatal density (SD: number of stomata per mm2 leaf area) and atmospheric concentration of CO2 ([CO2]) permits the use of plants as proxies of palaeo-atmospheric CO2. Many stomatal reconstructions of palaeo-[CO2] are based upon multiple fossil species. However, it is unclear how plants respond to [CO2] across genus, family or ecotype in terms of SD or stomatal index (SI: ratio of stomata to epidermal cells). This study analysed the stomatal numbers of conifers from the ancient family Cupressaceae, in order to examine the nature of the SI–[CO2] relationship, and potential implications for stomatal reconstructions of palaeo-[CO2].

Methods

Stomatal frequency measurements were taken from historical herbarium specimens of Athrotaxis cupressoides, Tetraclinis articulata and four Callitris species, and live A. cupressoides grown under CO2-enrichment (370, 470, 570 and 670 p.p.m. CO2).

Key Results

T. articulata, C. columnaris and C. rhomboidea displayed significant reductions in SI with rising [CO2]; by contrast, A. cupressoides, C. preissii and C. oblonga show no response in SI. However, A. cupressoides does reduce SI to increases in [CO2] above current ambient (approx. 380 p.p.m. CO2). This dataset suggests that a shared consistent SI–[CO2] relationship is not apparent across the genus Callitris.

Conclusions

The present findings suggest that it is not possible to generalize how conifer species respond to fluctuations in [CO2] based upon taxonomic relatedness or habitat. This apparent lack of a consistent response, in conjunction with high variability in SI, indicates that reconstructions of absolute palaeo-[CO2] based at the genus level, or upon multiple species for discrete intervals of time are not as reliable as those based on a single or multiple temporally overlapping species.  相似文献   

11.
Simple sequence repeats (SSRs) in the NCBI dbEST database were surveyed to identify potential SSR markers for Quercus mongolica. In total, 2,691 gene sequences, mainly from expressed sequence tags (ESTs) for Q. robur and Q. petraea had been registered. Twenty-two PCR primers were designed for SSRs in these sequences and screened for polymorphisms in 16 Q. mongolica trees. Ten loci were easily genotyped and showed polymorphism, with numbers of alleles and expected heterozygosity ranging from 3 to 15 and 0.28 to 0.94, respectively. These EST-SSR markers should be useful for studying the genetic diversity of Quercus species.  相似文献   

12.
Allozyme variation at 14 loci was studied electrophoretically in 19 population samples of Quercus cerris, Q. pubescens, Q. petraea, and Q. robur. Genetic variation, measured as mean heterozygosity, was found to be high within populations but low among populations. A significant deficit of heterozygotes was observed in most of the populations of Q. pubescens, Q. petraea, and Q. robur. At the species level, Q. cerris (sect. Cerris) appears to be clearly separated from the rest (sect. Quercus). Q. pubescens, Q. petraea, and Q. robur form a cluster of closely-related species. The degree of genetic differentiation among these species is low, i.e. in the range normally found among con-specific populations. Nevertheless, allozyme data allow the elaboration of a dendrogram which is in reasonable accordance with the taxonomic classification, but also supports the importance of hybridization and introgression. This is also underlined by the fact that the allelic differentiation at the Got-2 locus corresponds with the phenotypic classification (petraea-like, hybrids, and robur-like) in samples from mixed populations.  相似文献   

13.
14.
Mediavilla  S.  Santiago  H.  Escudero  A. 《Photosynthetica》2002,40(4):553-559
In the evergreen Quercus rotundifolia and the co-existing deciduous Q. faginea we studied the diurnal variations in photosynthetic capacity (P max), measured as the rate of O2 evolution at photon and CO2 saturation, and in the rate of net CO2 assimilation (P N) in the field during the period of maximum photosynthetic activity. Our aim was to check the contribution of stomatal and non-stomatal limitations to the diurnal variation in photosynthesis, and to study the differences between both species. Q. faginea leaves displayed lower mass per unit area and higher nitrogen content than Q. rotundifolia leaves. The maximum stomatal conductance and P N in the field were higher in Q. faginea than in Q rotundifolia. Also P max of Q. faginea was higher than that of Q. rotundifolia. Both species attained in the field a high percentage of the P max (around 82 % for Q. faginea and 73 % for Q. rotundifolia). This indicates reduced stomatal limitation of photosynthesis under favourable conditions, especially in Q. faginea. P N underwent a sharp decrease towards mid-day in association with increase in the atmospheric vapour pressure deficit and decrease in the leaf water potential. P max was also reduced during mid-day. This demonstrated the contribution of mesophyll limitations to the P N in the two species under stress. The mesophyll limitation of photosynthesis seemed to be similar for both species, independently from the differences in leaf traits between them.  相似文献   

15.
M. B. Jones 《Oecologia》1987,71(3):355-359
Summary Photosynthesis and transpiration was measured in the large emergent C4 sedge Cyperus papyrus (papyrus) which occupies wide areas of wetland on the African continent. The maximum observed value of net assimilation was 35 mol CO2 m-2 s-1 at full sunlight but light saturation of photosynthesis did not occur. The quantum yield of photosynthesis obtained from the initial slope of the light response curves (0.06 mol mol-1 incident light) was relatively high and close to previously recorded values for some C4 grasses. Measurements made over two days showed that stomatal conductance was sensitive to the ambient air vapour pressure deficit (VPD) and was consistently lower on the day when VPD's were higher. There was, however, no marked midday closure of the stomata. Photosynthesis was also reduced on the day when VPD's were higher. The relationship between net photosynthesis and stomatal conductance was close to linear over the range of measurement conditions, with the result that intercellular CO2 concentrations (C i ) did not vary markedly. There was some evidence that C i decreased at high VPD's. The regulation of stomatal movement in papyrus appears to minimise excessive water loss while not severely limiting photosynthesis. The significance of this strategy for a wetland species with plentiful supplies of water is discussed.  相似文献   

16.
We examined how independent and interactive effects of CO2 concentrations, water supply and wind speed affect growth rates, biomass partitioning, water use efficiency, diffusive conductance and stomatal density of plants. To test the prediction that wind stress will be ameliorated by increased CO2 and/or by unrestricted water supply we grew Sinapis alba L. plants in controlled chambers under combinations of two levels of CO2 (350 ppmv, 700 ppmv), two water regimes and two wind speeds (0.3 ms–1, 3.7 ms–1). We harvested at ten different dates over a period of 60 days. A growth analysis was carried out to evaluate treatment effects on plant responses. Plants grown both in increased CO2 and in low wind conditions had significantly greater stem length, leaf area and dry weights of plant parts. Water supply significantly affected stem diameter, root weight and leaf area. CO2 enrichment significantly increased the rate of biomass accumulation and the relative ratio of biomass increase to leaf area expansion. High wind speed significantly reduced plant growth rates and the rate of leaf area expansion was reduced more than the rate of biomass accumulation. Regression analysis showed significant CO2 effects on the proportion of leaf and stem dry weight to total dry weight. A marked plant-age effect was dependent on water supply, wind speed and CO2 concentration. A reduced water supply significantly decreased the stomatal conductance, and water use efficiency significantly increased with a limited water supply, low wind and increased CO2. We found significant CO2 x wind effects for water diffusion resistance, adaxial number of stomata and water use efficiencies and significant wind x water effect for water use efficiency. In conclusion, wind stress was ameliorated by growing in unrestricted water but not by growing in increased CO2.  相似文献   

17.
Attached leaves of Zea mays were illuminated with monochromatic light, with either the upper or the lower epidermis facing the light source. The mesophyll absorbed between 99.5 and 99.6% of the red or blue light used. An inversion of the light direction therefore caused a 200- to 250-fold change in the quantum flux into each epidermis. This variation in quantum flux did not affect stomatal conductance. Stomatal conductance was however correlated with intercellular CO2 concentration, ci, and the relationship between stomatal conductance and ci appeared also to remain the same if changes in ci were brought about by changes in atmospheric CO2 concentration instead of light. A close inspection of the data showed that stomata of the upper (adaxial) epidermis exhibited a small increase in conductance (<0.1 cm s-1) in response to blue light that was superimposed on the dominating response to ci.  相似文献   

18.
Reproductive strategies of closely related species distributed along successional gradients should differ as a consequence of the trade‐off between competition and colonization abilities. We compared male reproductive strategies of Quercus robur and Q. petraea, two partly interfertile European oak species with different successional status. In the studied even‐aged stand, trees of the late‐successional species (Q. petraea) grew faster and suffered less from intertree competition than trees of the early‐successional species (Q. robur). A large‐scale paternity study and a spatially explicit individual‐based mating model were used to estimate parameters of pollen production and dispersal as well as sexual barriers between species. Male fecundity was found to be dependent both on a tree's circumference and on its environment, particularly so for Q. petraea. Pollen dispersal was greater and more isotropic in Q. robur than in Q. petraea. Premating barriers to hybridization were strong in both species, but more so in Q. petraea than in Q. robur. Hence, predictions based on the competition–colonization trade‐off are well supported, whereas the sexual barriers themselves seem to be shaped by colonization dynamics.  相似文献   

19.
Background and Aims Following the consensus view for unitary origin and conserved function of stomata across over 400 million years of land plant evolution, stomatal abundance has been widely used to reconstruct palaeo-atmospheric environments. However, the responsiveness of stomata in mosses and hornworts, the most basal stomate lineages of extant land plants, has received relatively little attention. This study aimed to redress this imbalance and provide the first direct evidence of bryophyte stomatal responsiveness to atmospheric CO2.Methods A selection of hornwort (Anthoceros punctatus, Phaeoceros laevis) and moss (Polytrichum juniperinum, Mnium hornum, Funaria hygrometrica) sporophytes with contrasting stomatal morphologies were grown under different atmospheric CO2 concentrations ([CO2]) representing both modern (440 p.p.m. CO2) and ancient (1500 p.p.m. CO2) atmospheres. Upon sporophyte maturation, stomata from each bryophyte species were imaged, measured and quantified.Key Results Densities and dimensions were unaffected by changes in [CO2], other than a slight increase in stomatal density in Funaria and abnormalities in Polytrichum stomata under elevated [CO2].Conclusions The changes to stomata in Funaria and Polytrichum are attributed to differential growth of the sporophytes rather than stomata-specific responses. The absence of responses to changes in [CO2] in bryophytes is in line with findings previously reported in other early lineages of vascular plants. These findings strengthen the hypothesis of an incremental acquisition of stomatal regulatory processes through land plant evolution and urge considerable caution in using stomatal densities as proxies for paleo-atmospheric CO2 concentrations.  相似文献   

20.
To characterise the stomata of six temperate deciduous tree species, sets of stomatal sensitivities to all the most important environmental factors were measured. To compare the importance of abscisic acid (ABA) in the different stomatal responses, the effect of exogenous ABA on all the stomatal sensitivities was determined.Almost all the stomatal sensitivities: the sensitivity to a decrease in leaf water potential, air humidity, CO2 concentration ([CO2]) and light intensity, and to an increase in [CO2] and light intensity were the highest in the slow-growing species, and the lowest in the fast-growing species. Drought increased the sensitivity to the environmental changes that induce a decrease in the stomatal conductance, and decreased the sensitivity to the changes that induce an increase in this conductance. The sensitivities of the slow-growers were most strongly affected by drought and ABA. Therefore the success of the slow-growers in their ecological niches can be based on the highly sensitive and strictly regulated responses of their stomata. The fast-growers had the highest sensitivity to an increase in leaf water potential and this sensitivity was sharply reduced by drought and ABA. Thus, the dominance of the trees in riparian areas can be based on the ability of their stomata to quickly reach high conductance in well-watered conditions and to efficiently decrease this rate during drought.Stomatal sensitivities to the hydraulic environmental factors (water potentials in plant and air) had higher values in well-watered trees and a more pronounced response to drought than the sensitivities to the photosynthetic environmental factors ([CO2] and light intensity). Thus, the hydraulic factors most likely prevail over the photosynthetic factors in determining stomatal conductance in these species.In response to exogenous ABA, the rates of stomatal closure, following a decrease in air humidity and light intensity, and an increase in [CO2], were accelerated. Stomatal opening following an increase in air humidity and light intensity and a decrease in [CO2] was replaced by slow closing. The rate of stomatal opening following an increase in leaf water potential was reduced. As the sensitivities to changes in light were modified less by the ABA than the other stomatal sensitivities, the prediction of stomatal responses on the basis of the sensitivity to light alone should be excluded in stomatal models.  相似文献   

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