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1.
Leaves from Paphiopedilum sp. (Orchidaceae) having achlorophyllous stomata, show reduced levels of stomatal conductance when irradiated with red light, as compared with either the related, chlorophyllous genus Phragmipedium or with their response to blue light. These reduced levels of stomatal conductance, and the failure of isolated Paphiopedilum stomata to open under red irradiation indicates that the small stomatal response measured in the intact leaf under red light is indirect.

The overall low levels of stomatal conductance observed in Paphiopedilum leaves under most growing conditions and their capacity to increase stomatal conductance in response to blue light suggested that growth and carbon gain in Paphiopedilum could be enhanced in a blue light-enriched environment. To test that hypothesis, plants of Paphiopedilum acmodontum were grown in controlled growth chambers under daylight fluorescent light, with or without blue light supplementation. Total photosynthetic photon flux density was kept constant in both conditions. Blue light enrichment resulted in significantly higher growth rates—of up to 77%—over a 3 to 4 week growing period, with all evidence indicating that the blue light effect was a stomatal response. Manipulations of stomatal properties aimed at long-term carbon gains could have agronomic applications.

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2.
When leaves of plants with C3 metabolism are detached and held in darkness, they senesce and the stomata close. Because the relation of senescence and stomatal closure is very close, if not actually causal, the question arose as to whether in the leaves of plants with Crassulacean acid metabolism whose stomata open at night the relationship to senescence would be reversed. Detached leaves of four species of Hoya, floated on water in constant darkness or constant light, were found to show no large differences in stomatal aperture (measured as diffusion resistance) between those in the light or dark, but the aperture changed in a regular circadian rhythm. In some leaves the rhythm was simple, in others the peak showed small secondary peaks, but in all cases the values were nearly the same in the light as in the dark, throughout the cycle. Previous culture of the intact plants under normal day/night conditions gave results similar to those with plants that had had prolonged culture under constant light or darkness. In those cases when the stomata were more open in the dark, the chlorophyll content was greater than when the stomata were more open in the light; but when they were more open in the light, the chlorophyll content showed little difference between light and dark. When the leaves had only their petioles in water they showed greater senescence in the light than in the dark, and the stomata were more tightly closed in the light, especially at the apical ends. All four species of Hoya gave similar results. We deduce that senescence of these leaves is modified by stomatal aperture, and generally in the same direction as in C3 leaves, but that in continuous light or darkness the primary control over the aperture is the endogenous cycle.  相似文献   

3.
Pisum sativum L. cv Alsweet (garden pea) and Lycopersicon esculentum flacca Mill. (tomato) were used to evaluate the phytotoxicity of SO2 and O3 in the light and dark. Plants were grown in controlled environment chambers and exposed to SO2 or O3 in the light or dark at the same environmental conditions at which they were grown. The pea plants were treated with fusicoccin to ensure open stomata in the dark; the stomata of the tomato mutant remained open in the dark. Both species exhibited 64% to 80% less foliar necrosis following exposure to SO2 (0.5 to 1.0 microliter per liter for 2 hours) in the light than in the dark. The decrease in SO2 injury for light versus dark exposed plants was greater in fully expanded than expanding leaves. Both species exhibited 30% greater foliar necrosis following exposure to O3 (0.2 microliter per liter for 2 hours) in the light than dark. The increase in O3 injury in the light versus dark was similar for leaves at all stages of expansion. Leaf conductance to water vapor was 7% to 11% and 23% higher in the light than dark for fusicoccin-treated peas and tomato plants, respectively, indicating greater foliar uptake of both pollutants in the light than dark. Thus, the decreased SO2 toxicity in the light was not associated with pollutant uptake, but rather the metabolism of SO2. In contrast, the increased toxicity of O3 in the light was at least in part associated with increased uptake or could not be separated from it.  相似文献   

4.
Stomata usually open when leaves are transferred from darkness to light. However, reverse-phase stomatal opening in succulent plants has been known. CAM plants such as cacti and Opuntia ficus–indica achieve their high water use efficiency by opening their stomata during the cool, desert nights and closing them during the hot, dry days. Signal transduction pathway for stomatal opening by blue light photoreceptors including phototropins and the carotenoid pigment zeaxanthin has been suggested. Blue light regulated signal transduction pathway on stomatal opening could not be applied to CAM plants, but the most possible theory for a nocturnal response of stomata in CAM plants is photoperiodic circadian rhythm.  相似文献   

5.
Blue light induced stomatal opening has been studied by applying a short pulse (~5 to 60 s) of blue light to a background of saturating photosynthetic red photons, but little is known about steady-state stomatal responses. Here we report stomatal responses to blue light at high and low CO2 concentrations. Steady-state stomatal conductance (gs) of C3 plants increased asymptotically with increasing blue light to a maximum at 20% blue (120 μmol m−2 s−1). This response was consistent from 200 to 800 μmol mol−1 atmospheric CO2 (Ca). In contrast, blue light induced only a transient stomatal opening (~5 min) in C4 species above a Ca of 400 μmol mol−1. Steady-state gs of C4 plants generally decreased with increasing blue intensity. The net photosynthetic rate of all species decreased above 20% blue because blue photons have lower quantum yield (moles carbon fixed per mole photons absorbed) than red photons. Our findings indicate that photosynthesis, rather than a blue light signal, plays a dominant role in stomatal regulation in C4 species. Additionally, we found that blue light affected only stomata on the illuminated side of the leaf. Contrary to widely held belief, the blue light-induced stomatal opening minimally enhanced photosynthesis and consistently decreased water use efficiency.  相似文献   

6.
Alterations in light quality affect plant morphogenesis and photosynthetic responses but the effects vary significantly between species. Roses exhibit an irradiance‐dependent flowering control but knowledge on light quality responses is scarce. In this study we analyzed, the responses in morphology, photosynthesis and flowering of Rosa × hybrida to different blue (B) light proportions provided by light‐emitting diodes (LED, high B 20%) and high pressure sodium (HPS, low B 5%) lamps. There was a strong morphological and growth effect of the light sources but no significant difference in total dry matter production and flowering. HPS‐grown plants had significantly higher leaf area and plant height, yet a higher dry weight proportion was allocated to leaves than stems under LED. LED plants showed 20% higher photosynthetic capacity (Amax) and higher levels of soluble carbohydrates. The increase in Amax correlated with an increase in leaf mass per unit leaf area, higher stomata conductance and CO2 exchange, total chlorophyll (Chl) content per area and Chl a/b ratio. LED‐grown leaves also displayed a more sun‐type leaf anatomy with more and longer palisade cells and a higher stomata frequency. Although floral initiation occurred at a higher leaf number in LED, the time to open flowers was the same under both light conditions. Thereby the study shows that a higher portion of B light is efficient in increasing photosynthesis performance per unit leaf area, enhancing growth and morphological changes in roses but does not affect the total Dry Matter (DM) production or time to open flower.  相似文献   

7.
To learn how species differences in stomatal behavior are regulated, the response of epidermal and leaf diffusive resistance to light was investigated in Lycopersicon esculentum Mill., Solanum pennellii Corr., and a periclinal chimera having an S. pennellii epidermis and an L. esculentum mesophyll that was produced from a graft of the two species. S. pennellii has about 23% fewer stomata per square millimeter than does L. esculentum, and the two species have contrasting stomatal sensitivities to light. The abaxial stomata of L. esculentum open in dimmer light and to a greater extent than the adaxial stomata. The abaxial and adaxial stomata of S. pennellii respond similarly to light incident on the adaxial epidermis and are less open at all quantum flux densities than comparable stomata of L. esculentum. The patterns of response to light of the abaxial and adaxial stomata of the chimera were practically identical to those of L. esculentum, and quite unlike those of S. pennellii. Thus, the pattern of stomatal light response in the chimera was regulated by the L. esculentum mesophyll. The reduction in stomatal frequency of the chimera, which was regulated by the epidermis of S. pennellii, contributed to the 40% difference in leaf diffusive resistance between the plants in moderate light.  相似文献   

8.
Opening and closure movements of individual stomata were analyzed by light microscopy, scanning electron microscopy, and time-lapse photomicrography. Turgor-pressure-induced changes in pore shape of Commelina communis L. stomata were observed in vivo and in fixed material. The ventral wall of the guard cells undergoes three-dimensional alterations during opening and closing. Stomatal aperture increases with increase in light intensity and with decrease in CO2 concentration as previously described, while reactions to relative humidity and anaerobiosis are somewhat divergent from common experience. Low humidity induces opening rather than closure of stomata in well watered greenhouse plants, and lack of oxygen induces consecutive opening and closure movements. Individual stomata characteristically open by three distinct steps: (1) rapid opening; (2) immobility or slow closure; (3) relatively rapid opening. Since the timing of the second step varies for individual stomata, it is obscured by methods that integrate the responses of several stomata. The movements of individual stomata are well synchronized, but active communication between stomata in different areas can not be confirmed.  相似文献   

9.
Physiological and isotopic aspects of photosynthesis in peperomia   总被引:2,自引:2,他引:0       下载免费PDF全文
Physiological and isotopic aspects of several Peperomia species were investigated. All but one species had C3-like stomatal behavior, in that stomata were open during the day and closed during the night. In these species, most atmospheric CO2 uptake occurred during the day. Concurrent with this stomatal behavior, there were Crassulacean acid metabolism-like acid fluctuations in most species. Carbon and hydrogen isotope ratios of cellulose nitrate from Peperomia reflect their physiological behavior. The δ13C values of cellulose nitrate from Peperomia species were similar to values observed in C3 plants and consistent with the daytime uptake of exogeneous CO2 via the C3 photosynthetic pathway. The δD values of cellulose nitrate from Peperomia species approach those of Crassulacean acid metabolism plants. These elevated δD values are caused by fractionations occurring during biochemical reactions and not as a consequence of water relations.  相似文献   

10.
The activity of NADP and O2 photoreduction by water is essentially higher in chloroplasts isolated from pea seedlings (Pisum sativum L.) grown under blue light as compared with that from plants grown under red light. In contrast, the photoreduction of NADP and O2 with photosystem I only is practically the same or even lower in chloroplasts isolated from plants grown under blue light. The addition of plastocyanin does not affect the rate or the extent of NADP photoreduction by water in the chloroplasts isolated from plants grown under blue light, whereas it sharply activates NADP reduction in the chloroplasts isolated from plants grown under red light. The extent of the light-induced oxidation of cytochrome f is appreciably higher in chloroplasts isolated from plants grown under blue light. Cytochrome b559 plays the predominant role in the oxidoreductive reactions of these chloroplasts. Furthermore, the fluorescence measurements indicate more effective transfer of excitation energy from chlorophyll to the photosystem II reaction center in chloroplasts isolated from plants grown under blue light.  相似文献   

11.
The effects of environmental parameters on the blue light response of stomata were studied by quantifying transient increases in stomatal conductance in Commelina communis following 15 seconds by 0.100 millimole per square meter per second pulses of blue light. Because conductance increases were not observed following red light pulses of the same or greater (30 seconds by 0.200 millimole per square meter per second) fluences, the responses observed could be reliably attributed to the specific blue light response of the guard cells, rather than to guard cell chlorophyll. In both Paphiopedilum harrisianum, which lacks guard cell chloroplasts, and Commelina, the blue light response was enhanced by 0.263 millimole per square meter per second continuous background red light. Thus, the blue light response and its enhancement do not require energy derived from red-light-driven photophosphorylation by the guard cell chloroplasts. In Commelina, reduction of the intercellular concentration of CO2 by manipulation of ambient CO2 concentrations resulted in an enhanced blue light response. In both Commelina and Paphiopedilum, the blue light response was decreased by an increased vapor pressure difference. The magnitude of blue-light-specific stomatal opening thus appears to be sensitive to environmental conditions that affect the carbon and water status of the plant.  相似文献   

12.
Summary Plant survival ex vitro requires functioning stomata, adequate cuticular wax composition and deposition, and normal morphological development. Light intensity, CO2 and relative humidity were altered inside an acoustic window mist reactor to study their effects on carnation (Dianthus caryophyllus) growth, stomata development, hyperhydration and epicuticular wax content. Increasing the light intensity from 65 to 145 μmol m−2 s−1 and enrichment of the gas phase with CO2 (1350 ppm) reduced the number of hyperhydrated plants from 75 to 25% and increased the percentage dry weight of normal (healthy) plants from 17 to 25%. Lowering the relative humidity (≈70% RH) surrounding the plants during the mist-off phase for the last 2 wk of culture reduced the number of hyperhydrated plants from 70 to 9% and also increased the percentage of dry weight of normal plants from 16 to 25%. The stomata on plants grown in conditions of high light or low humidity had smaller apertures and appeared sunken when compared to stomata from plants grown in low light and high relative humidity. The epicuticular wax profiles of plants from the greenhouse or Magenta boxes showed a distinct shift in wax compounds with developmental age, plant type (hyperhydrated or normal), and type of box that was used (vented or not). In addition, very different wax profiles were observed from plants grown in reactors with altered CO2 and light intensities.  相似文献   

13.
In open places, plants are exposed to higher fluence rates of photosynthetically active radiation and to higher red to far-red ratios than under the shade of neighbor plants. High fluence rates are known to increase stomata density. Here we show that high, compared to low, red to far-red ratios also increase stomata density in Arabidopsis (Arabidopsis thaliana). High red to far-red ratios increase the proportion of phytochrome B (phyB) in its active form and the phyB mutant exhibited a constitutively low stomata density. phyB increased the stomata index (the ratio between stomata and epidermal cells number) and the level of anphistomy (by increasing stomata density more intensively in the adaxial than in the abaxial face). phyB promoted the expression of FAMA and TOO MANY MOUTHS genes involved in the regulation of stomata development in young leaves. Increased stomata density resulted in increased transpiration per unit leaf area. However, phyB promoted photosynthesis rates only at high fluence rates of photosynthetically active radiation. In accordance to these observations, phyB reduced long-term water-use efficiency estimated by the analysis of isotopic discrimination against 13CO2. We propose a model where active phyB promotes stomata differentiation in open places, allowing plants to take advantage of the higher irradiances at the expense of a reduction of water-use efficiency, which is compensated by a reduced leaf area.Photosynthesis, transpiration, and transpiration efficiency, the ratio of carbon fixation to water loss, are key physiological traits considered by plant breeders when selecting productive and water-use efficient plants (Rebetzke et al., 2002; Richards, 2006; Passioura, 2007). Opening of the stomata allows the uptake of CO2 necessary for photosynthesis but it simultaneously increases the loss of water and the potential deterioration of the water status. Plants are finely tuned to efficiently face this dilemma. Under low levels of photosynthetically active radiation (PAR), stomata open just enough to prevent the limitation of photosynthesis by CO2 influx and the photochemical phase of photosynthesis is the limiting step. If PAR increases, allowing higher rates of photochemical reactions, which leads to more ATP and NADPH, stomatal conductance also increases to allow sufficient CO2 to use these products in the Calvin cycle (Donahue et al., 1997; Yu et al., 2004). If instead of following this response coordinated to photosynthetic rates, stomata opened maximally in response to low PAR, more CO2 than needed would be allowed to reach the chloroplast at the expense of unnecessary water loss.Canopy shade light is characterized not only by reduced PAR levels but also by a reduced proportion of red light (R) compared to far-red light (FR) caused by the selective absorption of visible light by photosynthetic pigments and the reflection and transmission of FR (Holmes and Smith, 1977a). This low R/FR ratio compared to unfiltered sunlight is perceived by phytochromes (Smith, 1982; Ballaré et al., 1987; Pigliucci and Schmitt, 1999), mainly phytochrome B (phyB; Yanovsky et al., 1995). In Arabidopsis (Arabidopsis thaliana), the high R/FR signals perceived by phyB decrease the length of the stem and petioles, cause a more prostrate position of the leaves, and promote branching and delay flowering, among other responses (Reed et al., 1993; Franklin and Whitelam, 2005).Transgenic plants of potato (Solanum tuberosum) expressing the PHYB gene of Arabidopsis show higher stomatal conductance, transpiration rates, and photosynthesis rates per unit leaf area than the wild type (Thiele et al., 1999; Boccalandro et al., 2003; Schittenhelm et al., 2004). Stomata density is unaffected, indicating that phyB enhances the aperture of the stomatal pore in these transgenic plants. Stomatal conductance is higher in Fuchsia magellanica plants exposed to R than to FR pulses at the end of the photoperiod (Aphalo et al., 1991). However, there are no general effects of R/FR treatments on the aperture of the stomatal pore. The stomata of Commelina communis (Roth-Bejerano, 1981) and of the orchid of the genus Paphiopedilum (Talbott et al., 2002) open in response to R and this effect is reversed by FR, indicating a control by phytochrome. Nevertheless, this FR reversal of the effect of R is absent in wild-type Arabidopsis (Talbott et al., 2003). In Phaseolus vulgaris, FR accelerates stomatal movements during dark to light (opening) and light to dark (closing) transitions and this effect is R reversible, but phytochrome status has no effects under constant conditions of light or darkness (Holmes and Klein, 1985). In the latter species, prolonged FR added to a white-light background promotes stomatal conductance but this effect cannot be ascribed to phytochrome (Holmes et al., 1986).In addition to this rapid adjustment of the CO2 and water vapor fluxes to daily fluctuations in light levels via the regulation of the stomatal pore aperture, plants acclimate to the prevailing PAR conditions by changing stomatal density (number of stomata per unit area) and stomatal index (the ratio between the number of stomata in a given area and the total number of stomata and other epidermal cells in that same area). Stomatal density and stomatal index are higher in plants grown in full sunlight at high levels of PAR than in plants grown in shade (Willmer and Fricker, 1996; Lake et al., 2001; Thomas et al., 2004; Casson and Gray, 2008). Mature leaves sense the environment (light intensity and CO2) and produce a systemic signal that regulates stomatal density and index in young leaves (Coupe et al., 2006). A change in CO2 concentrations or PAR levels affects photosynthesis and therefore it was suggested that a metabolic compound associated to this process (i.e. a sugar) may regulate stomatal development (Coupe et al., 2006). However, there is no correlation between photosynthetic rate and stomatal index in poplar (Populus spp.; Miyazawa et al., 2006) and transgenic anti-small subunit of Rubisco tobacco (Nicotiana tabacum) plants, show reduced photosynthesis and normal responses of stomatal density and stomatal index to PAR, suggesting that other photoreceptors could be involved in this regulation (Baroli et al., 2008).Here we demonstrate that high, compared to low, R/FR ratios perceived by phyB increase stomata density, stomata index, and amphistomy in the leaves of Arabidopsis. This behavior results in an enhanced photosynthetic rate at high PAR at the expense of reduced water-use efficiency.  相似文献   

14.
The effect of light quality on the composition, function and structure of the thylakoid membranes, as well as on the photosynthetic rates of intact fronds from Asplenium australasicum, a shade plant, grown in blue, white, or red light of equal intensity (50 microeinsteins per square meter per second) was investigated. When compared with those isolated from plants grown in white and blue light, thylakoids from plants grown in red light have higher chlorophyll a/chlorophyll b ratios and lower amounts of light-harvesting chlorophyll a/b-protein complexes than those grown in blue light. On a chlorophyll basis, there were higher levels of PSII reaction centers, cytochrome f and coupling factor activity in thylakoids from red light-grown ferns, but lower levels of PSI reaction centers and plastoquinone. The red light-grown ferns had a higher PSII/PSI reaction center ratio of 4.1 compared to 2.1 in blue light-grown ferns, and a larger apparent PSI unit size and a lower PSII unit size. The CO2 assimilation rates in fronds from red light-grown ferns were lower on a unit area or fresh weight basis, but higher on a chlorophyll basis, reflecting the higher levels of electron carriers and electron transport in the thylakoids.

The structure of thylakoids isolated from plants grown under the three light treatments was similar, with no significant differences in the number of thylakoids per granal stack or the ratio of appressed membrane length/nonappressed membrane length. The large freeze-fracture particles had the same size in the red-, blue-, and white-grown ferns, but there were some differences in their density. Light quality is an important factor in the regulation of the composition and function of thylakoid membranes, but the effects depend upon the plant species.

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15.
Sensitivity to light quality and pigment composition were analysed and compared in abaxial and adaxial stomata of Gossypium barbadense L. (Pima cotton). In most plants, abaxial (lower) stomatal conductances are higher than adaxial (upper) ones, and stomatal opening is more sensitive to blue light than to red. In greenhouse-grown Pima cotton, abaxial stomatal conductances were two to three times higher than adaxial ones. In contrast, adaxial stomatal conductances were 1·5 to two times higher than abaxial ones in leaves from growth chamber-grown plants. To establish whether light quality was a factor in the regulation of the relationship between abaxial and adaxial stomatal conductances, growth-chamber-grown plants were exposed to solar radiation outdoors and to increased red light in the growth chamber. In both cases, the ratios of adaxial to abaxial stomatal conductance reverted to those typical of greenhouse plants. We investigated the hypothesis that adaxial stomata are more sensitive to blue light and abaxial stomata are more sensitive to red light. Measurements of stomatal apertures in mechanically isolated epidermal peels from growth chamber and greenhouse plants showed that adaxial stomata opened more under blue light than under red light, while abaxial stomata had the opposite response. Using HPLC, we quantified the chlorophylls and carotenoids extracted from isolated adaxial and abaxial guard cells. All pigments analysed were more abundant in the adaxial than in the abaxial guard cells. Antheraxanthin and β-carotene contents were 2·3 times higher in adaxial than in abaxial guard cells, comparing with ad/ab ratios of 1·5–1·9 for the other pigments. We conclude that adaxial and abaxial stomata from Pima cotton have a differential sensitivity to light quality and their distinct responses are correlated with different pigment content.  相似文献   

16.
  • C4 plants supply concentrated CO2 to bundle sheath (BS) cells, improving photosynthetic efficiency by suppressing photorespiration. Mesophyll chloroplasts in C4 plants are redistributed toward the sides of the BS cells (aggregative movement) in response to environmental stresses under light. Although this chloroplast movement is common in C4 plants, the significance and mechanisms underlying the aggregative movement remain unknown.
  • Under environmental stresses, such as drought and salt, CO2 uptake from the atmosphere is suppressed by closing stomata to prevent water loss. We hypothesized that CO2 limitation may induce the chloroplast aggregative movement. In this study, the mesophyll chloroplast arrangement in a leaf of finger millet, an NAD-malic enzyme type C4 plant, was examined under different CO2 concentrations and light conditions.
  • CO2 limitation around the leaves promoted the aggregative movement, but the aggregative movement was not suppressed, even at the higher CO2 concentration than in the atmosphere, under high intensity blue light. In addition, mesophyll chloroplasts did not change their arrangement under darkness or red light.
  • From these results, it can be concluded that CO2 limitation is not a direct inducer of the aggregative movement but would be a promoting factor of the movement under high intensity blue light.
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17.
Akita S  Moss DN 《Plant physiology》1973,52(6):601-603
Stomata of barley (Hordeum vulgare L.) and wheat (Triticum aestivum L.) leaves failed to open in the light and close in the dark or respond to changes in the CO2 concentration of the atmosphere in either light or dark when the leaves were in an O2-free atmosphere. In contrast, the expected responses to environmental changes were found in atmospheres containing 1.5% O2. It appears that O2 is necessary for both opening and closing of wheat and barley stomata.  相似文献   

18.
In Arabidopsis thaliana, the cryptochrome (CRY) blue light photoreceptors and the phytochrome (phy) red/far-red light photoreceptors mediate a variety of light responses. COP1, a RING motif–containing E3 ubiquitin ligase, acts as a key repressor of photomorphogenesis. Production of stomata, which mediate gas and water vapor exchange between plants and their environment, is regulated by light and involves phyB and COP1. Here, we show that, in the loss-of-function mutants of CRY and phyB, stomatal development is inhibited under blue and red light, respectively. In the loss-of-function mutant of phyA, stomata are barely developed under far-red light. Strikingly, in the loss-of-function mutant of either COP1 or YDA, a mitogen-activated protein kinase kinase kinase, mature stomata are developed constitutively and produced in clusters in both light and darkness. CRY, phyA, and phyB act additively to promote stomatal development. COP1 acts genetically downstream of CRY, phyA, and phyB and in parallel with the leucine-rich repeat receptor-like protein TOO MANY MOUTHS but upstream of YDA and the three basic helix-loop-helix proteins SPEECHLESS, MUTE, and FAMA, respectively. These findings suggest that light-controlled stomatal development is likely mediated through a crosstalk between the cryptochrome-phytochrome-COP1 signaling system and the mitogen-activated protein kinase signaling pathway.  相似文献   

19.
Crassulacean acid metabolism (CAM) was investigated in leaves and stems of the succulent C4 dicot Portulaca oleracea L. Diurnal acid fluctuations, CO2 gas exchange, and leaf resistance were monitored under various photoperiod and watering regimes. No CAM activity was seen in well watered plants grown under 16-hour days. Under 8-hour days, however, well watered plants showed a CAM-like pattern of acid fluctuation with amplitudes of 102 and 90 microequivalents per gram fresh weight for leaves and stems, respectively. Similar patterns were also observed in detached leaves and defoliated stems. Leaf resistance values indicated that stomata were open during part of the dark period, but night acidification most likely resulted from refixation of respiratory CO2. In water-stressed plants maximum acid accumulations were reduced under both long and short photoperiods. At night, these plants showed short periods of net CO2 uptake and stomatal opening which continued all night long during preliminary studies under natural environmental conditions. Greatest acid fluctuations, in P. oleracea, with amplitudes of 128 microequivalents per gram fresh weight, were observed in water-stressed plants which had been rewatered, especially when grown under short days. No net CO2 uptake took place, but stomata remained open throughout the night under these conditions. These results indicate that under certain conditions, such as water stress or short photoperiods, P. oleracea is capable of developing an acid metabolism with many similarities to CAM.  相似文献   

20.
Flooding the intercellular air spaces of leaves with water was shown to cause rapid closure of stomata in Tradescantia pallida, Lactuca serriola, Helianthus annuus, and Oenothera caespitosa. The response occurred when water was injected into the intercellular spaces, vacuum infiltrated into the intercellular spaces, or forced into the intercellular spaces by pressurizing the xylem. Injecting 50 mm KCl or silicone oil into the intercellular spaces also caused stomata to close, but the response was slower than with distilled water. Epidermis-mesophyll grafts for T. pallida were created by placing the epidermis of one leaf onto the exposed mesophyll of another leaf. Stomata in these grafts opened under light but closed rapidly when water was allowed to wick between epidermis and the mesophyll. When epidermis-mesophyll grafts were constructed with a thin hydrophobic filter between the mesophyll and epidermis stomata responded normally to light and CO2. These data, when taken together, suggest that the effect of water on stomata is caused partly by dilution of K+ in the guard cell and partly by the existence of a vapor-phase signal that originates in the mesophyll and causes stomata to open in the light.Stomatal responses to the environment have been studied in leaves for well over 100 years. More recently, the mechanisms for these responses have been investigated using isolated epidermes or isolated guard cell protoplasts. Despite the combination of these two approaches, the mechanisms by which stomata respond to environmental signals are not well understood. Since stomata control CO2 uptake and water loss from leaves, the responses of stomata to environmental factors are important determinants of terrestrial productivity and water use. It is therefore critical that we understand the mechanisms by which stomata respond to the environment if we are to accurately predict the effects of future climates on productivity and water cycles (Randall et al., 1996).There are two assumptions about stomata that are implicit in much of the recent literature: (1) that stomatal responses result from sensory mechanisms that reside within the guard cells, and (2) that stomata in isolated epidermes respond similarly to those in a leaf. The exception to this generalization is the stomatal response to humidity, which has been suggested to be the result of changes in guard cell water potential (Dewar, 1995, 2002) or of signaling from other cells in the leaf to the guard cells (Buckley et al., 2003). The assumption that guard cells directly sense CO2 and light is largely based on data from isolated epidermes that show effects of light and CO2 on stomatal apertures. As pointed out by Mott (2009), however, stomatal responses to light and CO2 in isolated epidermes are generally much different from those observed in leaves; e.g. responses in isolated epidermes are generally smaller than those in leaves, opening in response to light is slower, and closing in darkness is rarely observed. These observations were used to suggest that the mesophyll is somehow involved in stomatal responses to red light and CO2. This idea is supported by several recent studies that suggest that guard cells do not respond directly to red light. In the first of these studies it was shown that guard cells in an intact leaf do not show hyperpolarization of the plasma membrane in response to red light if the red light is applied to only the guard cell (Roelfsema et al., 2002). In contrast, blue light applied only to the guard cell does cause hyperpolarization, and red light does cause hyperpolarization if applied to the guard cell and the underlying mesophyll. The second study showed that stomata in albino areas of a leaf do not respond to red light, although they contain chloroplasts and do respond to blue light (Roelfsema et al., 2006). Finally, a third study has shown that isolated epidermes are much more sensitive to light and CO2 when placed in close contact with an exposed mesophyll from a leaf from the same or a different species (Mott et al., 2008). These epidermis-mesophyll grafts showed stomatal responses to light and CO2 that were indistinguishable from those in an intact leaf—a sharp contrast to the behavior of stomata in isolated epidermes that are floating on buffer solutions. In that study, illumination of a single stoma in a leaf using a small-diameter fiber optic did not produce stomatal opening, but opening did occur if several stomata and the underlying mesophyll were illuminated. Furthermore, this treatment actually caused opening of adjacent, but unilluminated, stomata (Mott et al., 2008).In constructing the epidermis-mesophyll grafts in the study described above (Mott et al., 2008), it was noticed that functional grafts could be produced only if both the mesophyll and the epidermis were blotted completely dry of any free water before placing them together. Although the tissues were apparently still fully hydrated, there was very little free water present (i.e. water not contained within the walls of the leaf cells), and both the mesophyll and epidermis felt and looked dry prior to assembly. In addition, even when free water was blotted away initially, stomata did not open in grafts that ended up with visible water on the epidermis or mesophyll that was caused by condensation during the experiment. These observations suggest that the presence of free water somehow prevented the stomata in the grafts from opening. Assuming that the mechanisms operating in the grafts were similar to those in an intact leaf, this result also suggests that free water may have an effect on stomata in leaves as well. In addition, it seems possible that the effect of free water on stomata could be related to the disruption of the signal from the mesophyll that was proposed in an earlier study (Mott et al., 2008). We hypothesize that disruption of this signal could be caused by (1) dilution of some solute that is necessary for opening (such as K+) in the guard cell walls, (2) dilution of an apoplastic, liquid-phase opening signal from the mesophyll to the guard cells, and (3) blockage of a vapor-phase opening signal from the mesophyll to the guard cells. This study was initiated to test these three hypotheses by examining the effect of free water and other liquids on stomatal functioning.  相似文献   

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