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1.
A comparison of the effects of a rapidly imposed water deficit with different leaf ages on chlorophyll a fluorescence and gas exchange was performed in maize (Zea mays L.) plants. The relationships between photosynthesis and leaf relative turgidity (RT) and ion leakage were further investigated. Leaf dehydration substantially decreased net photosynthetic rate (A) and stomatal conductance (G s), particularly for older leaves. With dehydration time, F v /F m maintained a relatively stable level for youngest leaves but significantly decreased for the older leaves. The electron transport rate (ETR) sharply decreased with intensifying dehydration and remained at lower levels during continuous dehydration. The photochemical quenching of variable chlorophyll fluorescence (q P) gradually decreased with dehydration intensity for the older leaves but increased for the youngest leaves, whereas dehydration did not affect the nonphotochemical chlorophyll fluorescence quenching (NPQ) for the youngest leaves but remarkably decreased it for the older leaves. The leaf RT was significantly and positively correlated with its F v /F m, ETR, and q P, and the leaf ion leakage was significantly and negatively correlated with F v /F m and NPQ. Our results suggest that the photosynthetic systems of young and old leaves decline at different rates when exposed to rapid dehydration.  相似文献   

2.
Cyanobacterium Nostoc commune is a species highly resistant against desiccation. In this study, we investigated changes in photochemical processes of photosynthesis and spectral reflectance indices during controlled desiccation of the colonies from Antarctica. In a dehydration process, water potential (WP) reached ?3 MPa and values of potential (F v/F m) and effective quantum yields (ΦPSII) of photosystem II were kept to high value until 90% of water was lost from the colony, and these values decreased rapidly by further loss of water. This indicates that the colony loses water mostly from the exopolysaccharidic envelope, not from cells during the initial part of dehydration (relative water content, RWC = 100–10%). Other suggestions of inhibition of photosynthetic processes after 90% loss of water were the increase of the chlorophyll fluorescence parameter F p/F s. The F m′ was higher than F m in hydrated colonies because of state transition which change energy distribution between PS I and PS II, but decreased to same level as F m in dehydrated colonies. The Normalized Difference Vegetation Index (NDVI) and Photochemical Reflectance Index (PRI) showed concave‐ and convex‐curvilinear relationship with RWC, respectively. The changes of NDVI values were, however, statistically insignificant. PRI values were predominantly below 0 because of phycobiliprotein involvement. These results were compared with the same species in the Arctic region. This is, according to our best knowledge, the first measurement of changes in spectral reflectance indices during desiccation of cyanobacteria.  相似文献   

3.
Abscisic acid (ABA) content and relative water content (RWC) in second fully expanded leaves of cold hardened plants and in dehydrated leaves of freezing tolerant barley (Hordeum vulgare L. cv. Lunet) were compared. ABA content and RWC in leaves did not change during the first day of cold hardening. On the contrary, dehydration of leaves led to a decrease of RWC and to an increase of ABA content.  相似文献   

4.

Three prevalent aliphatic polyamines (PAs) include putrescine, spermidine, and spermine; they are low-molecular-mass polycations involved in many physiological processes in plants, especially, under stressful conditions. In this experiment, three bean (Phaseolus vulgaris L.) genotypes were subjected to well-watered conditions and two moderate and severe water-stressed conditions with and without spermidine foliar application. Water stress reduced leaf relative water content (RWC), chlorophyll contents, stomatal conductance (gs), intercellular CO2 concentration (Ci), transpiration rate, maximal quantum yield of PSII (Fv/Fm), net photosynthetic rate (PN), and finally grain yield of bean plants. However, spermidine application elevated RWC, gs, Ci, Fv/Fm, and PN, which caused an increase in the grain yield and harvest index of bean plants under water stress. Overall, exogenous spermidine could be utilized to alleviate water stress through protection of photosynthetic pigments, increase of proline and carotenoid contents, and reduction of malondialdehyde content.

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5.
Water stress effects were studied on three cultivars ofEragrostis curvula. Leaf water potential, RWC, total plantleaf area, green dry weight mass percentage and CO2 gas-exchangeweremeasured during the onset of stress and after recovery. After 3 days of waterstress, RWC of cv Tanganyika plants was around 30–40% of controls,while RWC of cvs Ermelo and Consol was around 50–60% of controls.However midday and predawn water potentials were lower in cvs Tanganyka andErmelo than in cv Consol. After re-watering, RWC and water potentials recoveredonly in Consol plants. A strong decrease of leaf area was recorded in cvsErmeloand Consol during water stress (about 91–94% less than the leafarea of controls). Photosynthesis decreased as a function of the degree ofwaterstress severity in all cultivars. Also, light saturated photosynthesis,CO2 quantum yield and light at which saturated photosynthesisoccurred, were strongly reduced by water stress. Recovery of photosynthesis wasfound in cv Consol after five days re-watering. Cv Consol showed a betterconservation of water and higher resistance to water stress than the other twocvs.  相似文献   

6.
The changes in photosynthetic efficiency and photosynthetic pigments during dehydration of the resurrection plantSelaginella lepidophylla (from the Chiuhahuan desert, S.W. Texas, USA) were examined under different light conditions. Changes in the photosynthetic efficiency were deduced from chlorophyll a fluorescence measurements (Fo, Fm, and Fv) and pigment changes were measured by HPLC analysis. A small decrease in Fv/Fm was seen in hydrated stems in high light (650 μmol photons·m−2·s−1) but not in low light (50 μmol photons·m−2·s−1). However, a pronounced decline in Fv/Fm was observed during dehydration in both light treatments, after one to two hours of dehydration. A rise in Fo was observed only after six to ten hours of dehydration. Concomitant with the decrease in photosynthetic efficiency during dehydration a rise in the xanthophyll zeaxanthin was observed, even in low-light treatments. The increase in zeaxanthin can be related to previously observed photoprotective non-photochemical quenching of fluorescence in dehydrating stems ofS. lepidophylla. We hypothesize that under dehydrating conditions even low light levels become excessive and zeaxanthin-related photoprotection is engaged. We speculate that these processes, as well as stem curling and self shading (Eickmeier et al. 1992), serve to minimize photoinhibitory damage toS. lepidophylla during the process of dehydration.  相似文献   

7.
The effect of temperature, light-spectrum, desiccation and salinity gradients on the photosynthesis of a Japanese subtidal brown alga, Sargassum macrocarpum (Fucales), was determined using a pulse amplitude modulation-chlorophyll fluorometer and dissolved oxygen sensors. Temperature responses of the maximum (Fv/Fm in darkness) and effective (ΔF/Fm at 50 μmol photons m−2 s−1; = ΦPSII) quantum yields during 6-day culture (4–36°C) remained high at 12–28°C, but decreased at higher temperatures. Nevertheless, ΔF/Fm also dropped at temperatures below 8°C, suggesting light sensitivity under chilling temperatures because Fv/Fm remained high. Photosynthesis–irradiance responses at 24°C under red (660 nm), green (525 nm), blue (450 nm) and white light (metal halide lamp) showed that maximum net photosynthesis under blue and white light was greater than under red and green light, indicating the sensitivity and photosynthetic availability of blue light in the subtidal light environment. In the desiccation experiment, samples under aerial exposure of up to 8 h under dim-light at 24°C and 50% humidity showed that ΔF/Fm quickly declined after more than 45 min of emersion; furthermore, ΔF/Fm also failed to recover to initial levels even after 1 day of rehydration in seawater. Under the emersion state, the ΔF/Fm remained high when the relative water content (RWC) was greater than 50%; in contrast, it quickly dropped when the RWC was less than 50%. When the RWC was reduced below 50%, ΔF/Fm did not return to initial levels, regardless of subsequent re-hydration, suggesting a low capacity of photosynthesis to recover from desiccation. The stenohaline response of photosynthesis under 3-day culture is evident, given that ΔF/Fm declined when salinity was beyond 20–40 psu. Adaptation to subtidal environments in temperate waters of Japan can be linked to these traits.  相似文献   

8.
The possibility to improve the recovery of sugar beet plants after water stress by application of synthetic cytokinins N6-benzyladenine (BA) or N6-(m-hydroxybenzyl)adenosine (HBA) was tested. Relative water content (RWC), net photosynthetic rate (PN), transpiration rate (E), stomatal conductance (gs), chlorophyll (Chl) a and Chl b contents, and photosystem 2 efficiency characterized by variable to maximal fluorescence ratio (Fv/Fm) were measured in control plants, in water-stressed plants, and after rehydration (4, 8, 24, and 48 h). Water stress markedly decreased parameters of gas exchange, but they started to recover soon after irrigation. Application of BA or HBA to the substrate or sprayed on leaves only slightly stimulated recovery of PN, E, and gs in rehydrated plants, especially during the first phases of recovery. Chl contents decreased only under severe water stress and Fv/Fm ratio was not significantly affected by water stress applied. Positive effects of BA or HBA application on Chl content and Fv/Fm ratio were mostly not observed.  相似文献   

9.
Two species of eucalypt (Eucalyptus macrorhyncha and E. rossii) were grown under conditions of high temperatures (45 °C, maximum) and high light (1500 μmol m?2 s?1, maximum) at either ambient (350 μL L?1) or elevated (700 μL L?1) CO2 concentrations for 8 weeks. The growth enhancement, in terms of total dry weight, was 41% and 103% for E. macrorhyncha and E. rossii, respectively, when grown in elevated [CO2]. A reduction in specific leaf area and increased concentrations of non-structural carbohydrates were observed for leaves grown in elevated [CO2]. Plants grown in elevated [CO2] had an overall increase in photosynthetic CO2 assimilation rate of 27%; however, when measured at the same CO2 concentration a down-regulation of photosynthesis was evident especially for E. macrorhyncha. During the midday period when temperatures and irradiances were maximal, photosynthetic efficiency as measured by chlorophyll fluorescence (Fv/Fm) was lower in E. macrorhyncha than in E. rossii. Furthermore, Fv/Fm was lower in leaves of E. macrorhyncha grown under elevated than under ambient [CO2]. These reductions in Fv/Fm were accompanied by increases in both photochemical (qP) and nonphotochemical quenching (qN and NPQ), and by increases in the concentrations of xanthophyll cycle pigments with an increased proportion of the total xanthophyll cycle pool comprising of antheraxanthin and zeaxanthin. Thus, increased atmospheric [CO2] may enhance photoinhibition when environmental stresses such as high temperatures limit the capacity of a plant to respond with growth to elevated [CO2].  相似文献   

10.
To assess the vegetation status and substrate condition on seedling survival and flowery of an endangered plant species Aster kantoensis, I carried out an in situ seed-sowing experiment in the gravelly/sandy floodplain of the Kinu River, central Japan. The coverage of plant species, especially that of an alien species Eragrostis curvula, increased, and the subsequent reduction in the relative photosynthetically active photon flux density (PPFD) (%) after the large-scale flood in 1998 caused a decline in the percentage of seedling survival and flowery of A. kantoensis. Performances of A. kantoensis were compared under four types of seminatural conditions, i.e., combinations of existence or absence of E. curvula combined with gravelly or sandy substrates. The highest percentage of survival and flowery were observed on the gravelly substrate under the E. curvula,-free conditions, but values were low in areas invaded by E. curvula. Aggressive invasion of E. curvula was the principal cause of loss of safe sites for colonization of river-endemic plants.  相似文献   

11.
Changes in photochemical efficiency, non-radiative energy dissipation (NRD), de-epoxidation state of xanthophyll cycle components (DPS) and contents of the antioxidants ascorbic acid and glutathione were studied in leaves of the poikilohydric Ramonda serbica Panc. (Gesneriaceae) during cycles of dehydration and subsequent rehydration. In drying leaves, the intrinsic efficiency of PS II photochemistry and the photon yield of PS II electron transport showed strong progressive decreases. Simultaneously, the fraction of excitation energy dissipated as heat in the PS II antenna increased markedly. The energy-dependent component of non-photochemical quenching (NPQ) showed an increase in dehydrating leaves down to relative water contents (RWC) values near 30%. Further decreases in RWC below these values caused a decrease in NPQ. Accordingly, DPS showed a similar behaviour, with a sharp increase and a subsequent decrease at very low RWC, although the maximum DPS was reached at slightly lower RWC than that for the maximum NPQ. The pools of reduced ascorbate and glutathione increased strongly when the RWC values fell below 40% and remained high in fully dehydrated leaves. When plants were re-watered photosynthetic efficiency, NRD, DPS and antioxidant contents recovered their initial control values. However, during rehydration, the zeaxanthin content showed a transient increase, as did NPQ, indicating an increasing demand for non-radiative dissipation. On the other hand, the contents of reduced ascorbate and reduced glutathione decreased but were still relatively high in the initial phase of rehydration, when the rate of photosynthetic electron transport, proton pumping and NRD were still relatively low. These results indicate that several photoprotective mechanisms are operating in R. serbica. Protection from photo-oxidation and photoinhibition appears to be achieved by co-ordinated contributions by ascorbate, glutathione and zeaxanthin-mediated NPQ. This variety of photoprotective mechanisms may be essential for conferring desiccation-tolerance.This revised version was published online in October 2005 with corrections to the Cover Date.  相似文献   

12.
Abstract. Drought avoidance due to cuticular control increases with leaf number to a maximum in the intermediate leaves, decreasing to a minimum in the upper leaves. Dehydrated intermediate leaves do not rehydrate detectably when floated on water for several days. Excision of their petioles when submerged, permits full rehydration, presumably via the xylem. Stressing the plant by withholding water for 1–3 weeks fails to increase this already high resistance to water movement through the leaf surface. It does, however, markedly decrease the loss of water from the fully rehydrated (100% RWC) leaves during the first hour of dehydration, presumably due to a more rapid stomatal closure than in the non-stressed leaves. Dehydration tolerance increases with leaf number, without an intermediate maximum. The intermediate and upper leaves were markedly more tolerant of dehydration after drought-induced stress than when non-stressed. Dehydration tolerance in some cases, was inversely proportional to dehydration rate. It was possible, however, to equalize the rates of dehydration of drought-stressed and non-drought-stressed leaves without affecting the greater tolerance of the drought-stressed leaves. Dehydration avoidance by osmotic adjustment was markedly developed in the slowly dehydrated attached leaves of drought-stressed plants, but not in the rapidly dehydrated excised leaves. This is evidence of drought acclimation. It must, therefore, be concluded that the slow dehydration of the drought-stressed plants also leads to the increase in dehydration tolerance by permitting drought-induced acclimation. The overall drought resistance of cabbage leaves depends on the three components: drought avoidance, dehydration avoidance and dehydration tolerance. The latter two increase during acclimation but the cuticular control of drought avoidance does not.  相似文献   

13.
14.
Campuloclinium macrocephalum is an Asteraceous alien weed that invades roadside vegetation and grassland in South Africa. The role of allelopathy and competition in its invasiveness was investigated using Eragrostis curvula (weeping lovegrass, an indigenous grass), E. tef and Lactuca sativa (lettuce) as test species. Trials were conducted in Petri-dishes, pots and in the field. Root and shoot extracts of adult C. macrocephalum plants did not inhibit seed germination in any test species. The greatest effect was radicle stunting produced by leaf extracts at 10 and 25% w/v. Eragrostis curvula was less tolerant of the extracts than E. tef. Allelopathic effects could however not be confirmed in pot trials evaluating the interference potential of the weed or weed residue effects against E. curvula. E. curvula growth and biomass was not affected by plant densities of one or five C. macrocephalum per pot, whereas C. macrocephalum suffered a 17% mortality and density-dependant trade-offs of size and biomass for survival. Under field conditions C. macrocephalum had a broader ecological niche than E. curvula, invading hygrophilous and undisturbed grasslands not amenable for E. curvula establishment, this included well drained disturbed soils on which the latter proliferated. Evidence of competitive exclusion of E. curvula by C. macrocephalum or vice versa was not detected. The coexistence of both species irrespective of relative density suggested these species have different resource requirements. Allelopathy was not an adequate causal mechanism to explain invasiveness in Campuloclinium macrocephalum. A more traditional hypothesis such as the absence of natural enemies, at this stage, better justifies the weed’s invasion success.  相似文献   

15.
In this study, we investigated responses of the Photochemical Reflectance Index (PRI), and Normalized Difference Vegetation Index (NDVI) to gradual dehydration of several Antarctic lichen species (chlorolichens: Xanthoria elegans, Rhizoplaca melanophthalma, Physconia muscigena, cyanolichen: Leptogium puberulum), and a Nostoc commune colony from fully wet to a dry state. The gradual loss of physiological activity during dehydration was evaluated by chlorophyll fluorescence parameters. The experimental lichen species differed in thallus color, and intrathalline photobiont. In the species that did not exhibit color change with desiccation (X. elegans), NDVI and PRI were more or less constant (mean of 0.25, ??0.36, respectively) throughout a wide range of thallus hydration status showing a linear relation to relative water content (RWC). In contrast, the species with apparent species-specific color change during dehydration exhibited a curvilinear relation of NDVI and PRI to RWC. PRI decreased (R. melanophthalma, L. puberulum), increased (N. commune) or showed a polyphasic response (P. muscigena) with desiccation. Except for X. elegans, a curvilinear relation was found between the NDVI response to RWC in all species indicating the potential of combined ground research and remote sensing spectral data analyses in polar regions dominated by lichen flora. The chlorophyll fluorescence data recorded during dehydration (RWC decreased from 100 to 0%) revealed a polyphasic species-specific response of variable fluorescence measured at steady state—Fs, effective quantum yield of photosystem II (ΦPSII), and non-photochemical quenching (qN). Full hydration caused an inhibition of ΦPSII in N. commune while other species remained unaffected. The dehydration-dependent fall in ΦPSII was species-specific, starting at an RWC range of 22–32%. Critical RWC for ΦPSII was around 5–10%. Desiccation led to a species-specific polyphasic decrease in Fs and an increase in qN indicating the involvement of protective mechanisms in the chloroplastic apparatus of lichen photobionts and N. commune cells. In this study, the spectral reflectance and chlorophyll fluorescence data are discussed in relation to the potential of ecophysiological processes in Antarctic lichens, their resistance to desiccation and survival in Antarctic vegetation oases.  相似文献   

16.
The development of desiccation tolerance by vegetative tissues was an important step in the plants’ conquest of land. To counteract the oxidative stress generated under these conditions the xanthophyll cycle plays a key role. Recent reports have shown that desiccation itself induces de-epoxidation of xanthophyll cycle pigments, even in darkness. The aim of the present work was to study whether this trait is a common response of all desiccation-tolerant plants. The xanthophyll cycle activity and the maximal photochemical efficiency of PS II (F v/F m) as well as β-carotene and α-tocopherol contents were compared during slow and rapid desiccation and subsequent rehydration in six species pairs (with one desiccation-sensitive and one desiccation-tolerant species each) belonging to different taxa. Xanthophyll cycle pigments were de-epoxidised in darkness concomitantly with a decrease in F v/F m during slow dehydration in all the desiccation-tolerant species and in most of the desiccation-sensitive ones. De-epoxidation was reverted in darkness by re-watering in parallel with the recovery of the initial F v/F m. The stability of the β-carotene pool confirmed that its hydroxylation did not contribute to zeaxanthin formation. The α-tocopherol content of most of the species did not change during dehydration. Because it is a common mechanism present in all the desiccation-tolerant taxa and in some desiccation-sensitive species, and considering its role in antioxidant processes and in excess energy dissipation, the induction of the de-epoxidation of xanthophyll cycle pigments upon dehydration in the dark could be understood as a desiccation tolerance-related response maintained from the ancestral clades in the initial steps of land occupation by plants.  相似文献   

17.
Turgid pieces of mature maize roots were dried in air and progressive changes in their relative water content (RWC) determined. Viability was tested by reproducibility of the drying curves after dehydration to successively lower RWCs. After reaching a chosen RWC, the pieces were rehydrated (approximately 2 h), and a 2nd and 3rd dehydration curve measured. Each drying curve was characterized by two parameters (a scale parameter λ, and a shape parameter β) of a survivorship function, which is a linear function of time. The parameter λ is more informative, and does not change in successive dehydrations for RWC > 0·4, suggesting no irreversible damage to the roots. Damage and death were indicated by divergences of λ in successive dehydrations to RWC = 0·35–0·15. Cryo-analytical microscopy confirmed these data while indicating specifically death of 50 and 100% of cortical cells at RWC 0·30 and 0·15, respectively, and survival of 50% or more of sieve tubes, pericycle and vascular parenchyma cells at root RWC as low as 0·15. This pattern of stelar cell survival may allow roots to preserve their capacity for renewal of axial conductivity and branch root development following periods of severe water stress.  相似文献   

18.
Kalanchoë daigremontiana, a CAM plant grown in a greenhouse, was subjected to severe water stress. The changes in photosystem II (PSII) photochemistry were investigated in water‐stressed leaves. To separate water stress effects from photoinhibition, water stress was imposed at low irradiance (daily peak PFD 150 μmol m?2 s?1). There were no significant changes in the maximal efficiency of PSII photochemistry (Fv/Fm), the traditional fluorescence induction kinetics (OIP) and the polyphasic fluorescence induction kinetics (OJIP), suggesting that water stress had no direct effects on the primary PSII photochemistry in dark‐adapted leaves. However, PSII photochemistry in light‐adapted leaves was modified in water‐stressed plants. This was shown by the decrease in the actual PSII efficiency (ΦPSII), the efficiency of excitation energy capture by open PSII centres (Fv′/Fm′), and photochemical quenching (qP), as well as a significant increase in non‐photochemical quenching (NPQ) in particular at high PFDs. In addition, photoinhibition and the xanthophyll cycle were investigated in water‐stressed leaves when exposed to 50% full sunlight and full sunlight. At midday, water stress induced a substantial decrease in Fv/Fm which was reversible. Such a decrease was greater at higher irradiance. Similar results were observed in ΦPSII, qP, and Fv′/Fm′. On the other hand, water stress induced a significant increase in NPQ and the level of zeaxanthin via the de‐epoxidation of violaxanthin and their increases were greater at higher irradiance. The results suggest that water stress led to increased susceptibility to photoinhibition which was attributed to a photoprotective process but not to a photodamage process. Such a photoprotection was associated with the enhanced formation of zeaxanthin via de‐epoxidation of violaxanthin. The results also suggest that thermal dissipation of excess energy associated with the xanthophyll cycle may be an important adaptive mechanism to help protect the photosynthetic apparatus from photoinhibitory damage for CAM plants normally growing in arid and semi‐arid areas where they are subjected to a combination of water stress and high light.  相似文献   

19.
The current concentrations of O3 have been shown to cause significant negative effects on crop yield. The present levels of ozone may not induce visible symptoms in most of plants, but can result in substantial losses in reproductive output. This paper considers the impact of ambient O3 on gas exchange, photosynthetic pigments, chlorophyll (Chl) fluorescence and carbohydrate levels in the flag leaf of wheat plants during various stages of reproductive development using open-top chambers. Mean O3 concentration was 45.7 ppb during wheat growth and 50.2 ppb after flag leaf development. Reproductive stage showed higher exceedence of O3 above 40 ppb compared to the vegetative stage. Diurnal variations in net photosynthetic rate (P N) and stomatal conductance (g s), intercellular CO2 concentration (C i), Fv/Fm ratio, photosynthetic pigments, soluble sugars, and starch were measured at 10, 30, and 50 days after flag leaf expansion (DAFE). The results showed reductions in P N, g s, Fv/Fm ratio, photosynthetic pigments and starch, and increases in C i, F0, and soluble sugars in nonfiltered chambers (NFCs) compared to filtered chambers (FCs). Maximum changes in measured parameters were observed at 50 DAFE (i.e. grain filling and setting phase). Diurnal variation in P N showed double peaked curve in both FCs and NFCs, but delayed peak and early depression in NFCs. Stomatal conductance was significantly lower in NFCs. The study suggests that higher prevalence of ambient O3 during reproductive development led to significant alteration in physiological vitality of wheat having potential negative influence on yield.  相似文献   

20.
Photosynthetic responses to salinity in two clones of Agrostis stolonifera   总被引:2,自引:1,他引:1  
Abstract The response of photosynthesis, Fe, to salinity was examined in two clones of Agrostis stolonifera, one derived from a salt marsh (SM) and the other from an inland site (IL). The response of young and old leaves was examined over one week of salt treatment and changes in leaf water relations and concentrations of Na+, Cl? and K+ ions were also monitored. Sodium+ and chloride? concentrations rose much more sharply in the older leaves of both clones and Fe declined to ca. zero by Day 4. Fe in the young leaves of the SM clone was unaffected by salt whilst the young leaves of the IL clone showed a 20% reduction in Fe within 24 h of salt application. This decline was due to a decline in stomatal conductance and a reduction in the photosynthetic capacity of the mesophyll and was associated with a greater accumulation of Na+ and Cl? ions and loss of K+ ions than in the SM clone. The significance of the salt exclusion from the young leaves of the SM clone to its growth on the salt marsh is discussed.  相似文献   

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