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1.
Species are often classified along a continuum from isohydric to anisohydric, with isohydric species exhibiting tighter regulation of leaf water potential through stomatal closure in response to drought. We investigated plasticity in stomatal regulation in an isohydric (Eucalyptus camaldulensis) and an anisohydric (Acacia aptaneura) angiosperm species subject to repeated drying cycles. We also assessed foliar abscisic acid (ABA) content dynamics, aboveground/belowground biomass allocation and nonstructural carbohydrates. The anisohydric species exhibited large plasticity in the turgor loss point (ΨTLP), with plants subject to repeated drying exhibiting lower ΨTLP and correspondingly larger stomatal conductance at low water potential, compared to plants not previously exposed to drought. The anisohydric species exhibited a switch from ABA to water potential‐driven stomatal closure during drought, a response previously only reported for anisohydric gymnosperms. The isohydric species showed little osmotic adjustment, with no evidence of switching to water potential‐driven stomatal closure, but did exhibit increased root:shoot ratios. There were no differences in carbohydrate depletion between species. We conclude that a large range in ΨTLP and biphasic ABA dynamics are indicative of anisohydric species, and these traits are associated with exposure to low minimum foliar water potential, dense sapwood and large resistance to xylem embolism.  相似文献   

2.
Stomatal regulation is crucial for forest species performance and survival on drought‐prone sites. We investigated the regulation of root and shoot hydraulics in three Pinus radiata clones exposed to drought stress and its coordination with stomatal conductance (gs) and leaf water potential (Ψleaf). All clones experienced a substantial decrease in root‐specific root hydraulic conductance (Kroot‐r) in response to the water stress, but leaf‐specific shoot hydraulic conductance (Kshoot‐l) did not change in any of the clones. The reduction in Kroot‐r caused a decrease in leaf‐specific whole‐plant hydraulic conductance (Kplant‐l). Among clones, the larger the decrease in Kplant‐l, the more stomata closed in response to drought. Rewatering resulted in a quick recovery of Kroot‐r and gs. Our results demonstrated that the reduction in Kplant‐l, attributed to a down regulation of aquaporin activity in roots, was linked to the isohydric stomatal behaviour, resulting in a nearly constant Ψleaf as water stress started. We concluded that higher Kplant‐l is associated with water stress resistance by sustaining a less negative Ψleaf and delaying stomatal closure.  相似文献   

3.
Recent work has shown that stomatal conductance (gs) and assimilation (A) are responsive to changes in the hydraulic conductance of the soil to leaf pathway (KL), but no study has quantitatively described this relationship under controlled conditions where steady‐state flow is promoted. Under steady‐state conditions, the relationship between gs, water potential (Ψ) and KL can be assumed to follow the Ohm's law analogy for fluid flow. When boundary layer conductance is large relative to gs, the Ohm's law analogy leads to gs = KLsoilleaf)/D, where D is the vapour pressure deficit. Consequently, if stomata regulate Ψleaf and limit A, a reduction in KL will cause gs and A to decline. We evaluated the regulation of Ψleaf and A in response to changes in KL in well‐watered ponderosa pine seedlings (Pinus ponderosa). To vary KL, we systematically reduced stem hydraulic conductivity (k) using an air injection technique to induce cavitation while simultaneously measuring Ψleaf and canopy gas exchange in the laboratory under constant light and D. Short‐statured seedlings (< 1 m tall) and hour‐long equilibration times promoted steady‐state flow conditions. We found that Ψleaf remained constant near ? 1·5 MPa except at the extreme 99% reduction of k when Ψleaf fell to ? 2·1 MPa. Transpiration, gs, A and KL all declined with decreasing k (P < 0·001). As a result of the near homeostasis in bulk Ψleaf, gs and A were directly proportional to KL (R2 > 0·90), indicating that changes in KL may affect plant carbon gain.  相似文献   

4.
The water potential at turgor loss point (Ψtlp) has been suggested as a key functional trait for determining plant drought tolerance, because of its close relationship with stomatal closure. Ψtlp may indicate drought tolerance as plants, which maintain gas exchange at lower midday water potentials as soil water availability declines also have lower Ψtlp. We evaluated 17 species from seasonally dry habitats, representing a range of life‐forms, under well‐watered and drought conditions, to determine how Ψtlp relates to stomatal sensitivity (pre‐dawn water potential at stomatal closure: Ψgs0) and drought strategy (degree of isohydry or anisohydry; ΔΨMD between well‐watered conditions and stomatal closure). Although Ψgs0 was related to Ψtlp, Ψgs0 was better related to drought strategy (ΔΨMD). Drought avoiders (isohydric) closed stomata at water potentials higher than their Ψtlp; whereas, drought tolerant (anisohydric) species maintained stomatal conductance at lower water potentials than their Ψtlp and were more dehydration tolerant. There was no significant relationship between Ψtlp and ΔΨMD. While Ψtlp has been related to biome water availability, we found that Ψtlp did not relate strongly to stomatal closure or drought strategy, for either drought avoiders or tolerators. We therefore suggest caution in using Ψtlp to predict vulnerability to drought.  相似文献   

5.
We investigated how leaf hydraulic conductance (Kleaf) of loblolly pine trees is influenced by soil nitrogen amendment (N) in stands subjected to ambient or elevated CO2 concentrations (CO2a and CO2e, respectively). We also examined how Kleaf varies with changes in reference leaf water potential (Ψleaf‐ref) and stomatal conductance (gs‐ref) calculated at vapour pressure deficit, D of 1 kPa. We detected significant reductions in Kleaf caused by N and CO2e, but neither treatment affected pre‐dawn or midday Ψleaf. We also detected a significant CO2e‐induced reduction in gs‐ref and Ψleaf‐ref. Among treatments, the sensitivity of Kleaf to Ψleaf was directly related to a reference Kleaf (Kleaf‐ref computed at Ψleaf‐ref). This liquid‐phase response was reflected in a similar gas‐phase response, with gs sensitivity to D proportional to gs‐ref. Because leaves represented a substantial component of the whole‐tree conductance, reduction in Kleaf under CO2e affected whole‐tree water use by inducing a decline in gs‐ref. The consequences of the acclimation of leaves to the treatments were: (1) trees growing under CO2e controlled morning leaf water status less than CO2a trees resulting in a higher diurnal loss of Kleaf; (2) the effect of CO2e on gs‐ref was manifested only during times of high soil moisture.  相似文献   

6.
Relatively anisohydric species are predicted to be more predisposed to hydraulic failure than relatively isohydric species, as they operate with narrower hydraulic safety margins. We subjected co‐occurring anisohydric Juniperus monosperma and isohydric Pinus edulis trees to warming, reduced precipitation, or both, and measured their gas exchange and hydraulic responses. We found that reductions in stomatal conductance and assimilation by heat and drought were more frequent during relatively moist periods, but these effects were not exacerbated in the combined heat and drought treatment. Counter to expectations, both species exhibited similar gs temporal dynamics in response to drought. Further, whereas P. edulis exhibited chronic embolism, J. monosperma showed very little embolism due to its conservative stomatal regulation and maintenance of xylem water potential above the embolism entry point. This tight stomatal control and low levels of embolism experienced by juniper refuted the notion that very low water potentials during drought are associated with loose stomatal control and with the hypothesis that anisohydric species are more prone to hydraulic failure than isohydric species. Because direct association of stomatal behaviour with embolism resistance can be misleading, we advocate consideration of stomatal behaviour relative to embolism resistance for classifying species drought response strategies.  相似文献   

7.
The temperature dependence of mesophyll conductance (gm) was measured in well‐watered red raspberry (Rubus idaeus L.) plants acclimated to leaf‐to‐air vapour pressure deficit (VPDL) daytime differentials of contrasting amplitude, keeping a fixed diurnal leaf temperature (Tleaf) rise from 20 to 35 °C. Contrary to the great majority of gm temperature responses published to date, we found a pronounced reduction of gm with increasing Tleaf irrespective of leaf chamber O2 level and diurnal VPDL regime. Leaf hydraulic conductance was greatly enhanced during the warmer afternoon periods under both low (0.75 to 1.5 kPa) and high (0.75 to 3.5 kPa) diurnal VPDL regimes, unlike stomatal conductance (gs), which decreased in the afternoon. Consequently, the leaf water status remained largely isohydric throughout the day, and therefore cannot be evoked to explain the diurnal decrease of gm. However, the concerted diurnal reductions of gm and gs were well correlated with increases in leaf abscisic acid (ABA) content, thus suggesting that ABA can induce a significant depression of gm under favourable leaf water status. Our results challenge the view that the temperature dependence of gm can be explained solely from dynamic leaf anatomical adjustments and/or from the known thermodynamic properties of aqueous solutions and lipid membranes.?  相似文献   

8.
Risk-taking plants: Anisohydric behavior as a stress-resistance trait   总被引:1,自引:0,他引:1  
Water scarcity is a critical limitation for agricultural systems. Two different water management strategies have evolved in plants: an isohydric strategy and an anisohydric strategy. Isohydric plants maintain a constant midday leaf water potential (Ψleaf) when water is abundant, as well as under drought conditions, by reducing stomatal conductance as necessary to limit transpiration. Anisohydric plants have more variable Ψleaf and keep their stomata open and photosynthetic rates high for longer periods, even in the presence of decreasing leaf water potential. This risk-taking behavior of anisohydric plants might be beneficial when water is abundant, as well as under moderately stressful conditions. However, under conditions of intense drought, this behavior might endanger the plant. We will discuss the advantages and disadvantages of these two water-usage strategies and their effects on the plant’s ability to tolerate abiotic and biotic stress. The involvement of plant tonoplast AQPs in this process will also be discussed.  相似文献   

9.
We merge concepts from stomatal optimization theory and cohesion–tension theory to examine the dynamics of three mechanisms that are potentially limiting to leaf‐level gas exchange in trees during drought: (1) a ‘demand limitation’ driven by an assumption of optimal stomatal functioning; (2) ‘hydraulic limitation’ of water movement from the roots to the leaves; and (3) ‘non‐stomatal’ limitations imposed by declining leaf water status within the leaf. Model results suggest that species‐specific ‘economics’ of stomatal behaviour may play an important role in differentiating species along the continuum of isohydric to anisohydric behaviour; specifically, we show that non‐stomatal and demand limitations may reduce stomatal conductance and increase leaf water potential, promoting wide safety margins characteristic of isohydric species. We used model results to develop a diagnostic framework to identify the most likely limiting mechanism to stomatal functioning during drought and showed that many of those features were commonly observed in field observations of tree water use dynamics. Direct comparisons of modelled and measured stomatal conductance further indicated that non‐stomatal and demand limitations reproduced observed patterns of tree water use well for an isohydric species but that a hydraulic limitation likely applies in the case of an anisohydric species.  相似文献   

10.
The main factors regulating grapevine response to decreasing water availability were assessed under statistical support using published data related to leaf water relations in an extensive range of scion and rootstock genotypes. Matching leaf water potential (Ψleaf) and stomatal conductance (gs) data were collected from peer‐reviewed literature with associated information. The resulting database contained 718 data points from 26 different Vitis vinifera varieties investigated as scions, 15 non‐V. vinifera rootstock genotypes and 11 own‐rooted V. vinifera varieties. Linearised data were analysed using the univariate general linear model (GLM) with factorial design including biological (scion and rootstock genotypes), methodological and environmental (soil) fixed factors. The first GLM performed on the whole database explained 82.4% of the variability in data distribution having the rootstock genotype the greatest contribution to variability (19.1%) followed by the scion genotype (16.2%). A classification of scions and rootstocks according to their mean predicted gs in response to moderate water stress was generated. This model also revealed that gs data obtained using a porometer were in average 2.1 times higher than using an infra‐red gas analyser. The effect of soil water‐holding properties was evaluated in a second analysis on a restricted database and showed a scion‐dependant effect, which was dominant over rootstock effect, in predicting gs values. Overall the results suggest that a continuum exists in the range of stomatal sensitivities to water stress in V. vinifera, rather than an isohydric–anisohydric dichotomy, that is further enriched by the diversity of scion‐rootstock combinations and their interaction with different soils.  相似文献   

11.
Current understanding of physiological mechanisms governing stomatal behavior under water stress conditions is still incomplete and controversial. It has been proposed that coordination of stomatal kinetics with xylem vulnerability to cavitation [vulnerability curve (VC)] leads to different levels of isohydry/anisohydry in different plant species/cultivars. In this study, this hypothesis is tested in Vitis vinifera cultivars displaying contrasting stomatal behavior under drought stress. The cv Montepulciano (MP, near‐isohydric) and Sangiovese (SG, anisohydric) were compared in terms of stomatal response to leaf and stem water potential, as possibly correlated to different petiole hydraulic conductivity (kpetiole) and VC, as well as to leaf water relations parameters. MP leaves showed almost complete stomatal closure at higher leaf and stem water potentials than SG leaves. Moreover, MP petioles had higher maximum kpetiole and were more vulnerable to cavitation than SG. Water potential at the turgor loss point was higher in MP than in SG. In SG, the percentage reduction of stomatal conductance (PLgs) under water stress was almost linearly correlated with corresponding percentage loss of kpetiole (PLC), while in MP PLgs was less influenced by PLC. Our results suggest that V. vinifera near‐isohydric and anisohydric genotypes differ in terms of xylem vulnerability to cavitation as well as in terms of kpetiole, and that the coordination of these traits leads to their different stomatal responses under water stress conditions.  相似文献   

12.
Diurnal depression of leaf hydraulic conductance in a tropical tree species   总被引:10,自引:2,他引:8  
Diurnal patterns of hydraulic conductance of the leaf lamina (Kleaf) were monitored in a field‐grown tropical tree species in an attempt to ascertain whether the dynamics of stomatal conductance (gs) and CO2 uptake (Aleaf) were associated with short‐term changes in Kleaf. On days of high evaporative demand mid‐day depression of Kleaf to between 40 and 50% of pre‐dawn values was followed by a rapid recovery after 1500 h. Leaf water potential during the recovery stage was less than ?1 MPa implying a refilling mechanism, or that loss of Kleaf was not linked to cavitation. Laboratory measurement of the response of Kleaf to Ψleaf confirmed that leaves in the field were operating at water potentials within the depressed region of the leaf ‘vulnerability curve’. Diurnal courses of Kleaf and Ψleaf predicted from measured transpiration, xylem water potential and the Kleaf vulnerability function, yielded good agreement with observed trends in both leaf parameters. Close correlation between depression of Kleaf, gs and Aleaf suggests that xylem dysfunction in the leaf may lead to mid‐day depression of gas exchange in this species.  相似文献   

13.
Stomatal conductance (gs) and mesophyll conductance (gm) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (Kleaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, Kleaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H2O and CO2 diffusion inside leaves under varying light conditions. Gas exchange, Kleaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, gs, gm, and Kleaf were strongly correlated across species. However, the response patterns of A, gs, gm, and Kleaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.  相似文献   

14.
Drought stress is known to limit photosynthesis rates and to inflict photo‐oxidative damage in grapevines. Grapevines, which are considered drought‐tolerant plants, are characterized by diverse hydraulic and photosynthetic behaviors, depending on the cultivar. This research compared the photosynthesis and the photorespiration of Cabernet Sauvignon (Cs) (isohydric) and Shiraz (anisohydric) in an attempt to acquire a wider perspective on the iso/anisohydric phenomenon and its implications. Shiraz and Cs were subjected to terminal drought in the greenhouse. Soil water content (θ), leaf water potential (Ψl) and stomata conductance (gs) were measured to determine the cultivars' hydraulic behavior. Gas exchange and fluorometry measurements were taken at 21 and 2% O2 to acquire photosynthesis and photorespiration characteristics. Cs was found to behave in a near isohydric manner whereas Shiraz behaved in a near anisohydric manner. Compared to Shiraz, the reduced stomata conductance values of Cs were accompanied by higher water use efficiency and photorespiration rates, as well as photosystem II photochemical potential (Fv/Fm). As compared with Shiraz, Cs compensated for lower stomata conductance by higher photosynthesis and photorespiration. These two processes contributed to higher electron flow rates that might have a role in photoinhibition avoidance, which was observed in the stability of Fv/Fm under drought stress.  相似文献   

15.
Leaf gas‐exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas‐exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon–juniper Pinus edulisJuniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (?45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas‐exchange rates under well‐watered conditions, leaf‐specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade‐off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.  相似文献   

16.
Previous studies have reported correlation of leaf hydraulic vulnerability with pressure–volume parameters related to cell turgor. This link has been explained on the basis of the effects of turgor on connectivity among cells and tissue structural integrity, which affect leaf water transport. In this study, we tested the hypothesis that osmotic adjustment to water stress would shift the leaf vulnerability curve toward more negative water potential (Ψleaf) by increasing turgor at low Ψleaf. We measured leaf hydraulic conductance (Kleaf), Kleaf vulnerability [50 and 80% loss of Kleaf (P50 and P80); |Ψleaf| at 50 and 80% loss of Kleaf, respectively), bulk leaf water relations, leaf gas exchange and sap flow in two Vitis vinifera cultivars (Tempranillo and Grenache), under two water treatments. We found that P50, P80 and maximum Kleaf decreased seasonally by more than 20% in both cultivars and watering treatments. However, Kleaf at ?2 MPa increased threefold, while osmotic potential at full turgor and turgor loss point decreased. Our results indicate that leaf resistance to hydraulic dysfunction is seasonally plastic, and this plasticity may be mediated by osmotic adjustment.  相似文献   

17.
Identifying the drivers of stomatal closure and leaf damage during stress in grasses is a critical prerequisite for understanding crop resilience. Here, we investigated whether changes in stomatal conductance (gs) during dehydration were associated with changes in leaf hydraulic conductance (Kleaf), xylem cavitation, xylem collapse, and leaf cell turgor in wheat (Triticum aestivum). During soil dehydration, the decline of gs was concomitant with declining Kleaf under mild water stress. This early decline of leaf hydraulic conductance was not driven by cavitation, as the first cavitation events in leaf and stem were detected well after Kleaf had declined. Xylem vessel deformation could only account for <5% of the observed decline in leaf hydraulic conductance during dehydration. Thus, we concluded that changes in the hydraulic conductance of tissues outside the xylem were responsible for the majority of Kleaf decline during leaf dehydration in wheat. However, the contribution of leaf resistance to whole plant resistance was less than other tissues (<35% of whole plant resistance), and this proportion remained constant as plants dehydrated, indicating that Kleaf decline during water stress was not a major driver of stomatal closure.  相似文献   

18.
Stomata regulate CO2 uptake for photosynthesis and water loss through transpiration. The approaches used to represent stomatal conductance (gs) in models vary. In particular, current understanding of drivers of the variation in a key parameter in those models, the slope parameter (i.e. a measure of intrinsic plant water‐use‐efficiency), is still limited, particularly in the tropics. Here we collected diurnal measurements of leaf gas exchange and leaf water potential (Ψleaf), and a suite of plant traits from the upper canopy of 15 tropical trees in two contrasting Panamanian forests throughout the dry season of the 2016 El Niño. The plant traits included wood density, leaf‐mass‐per‐area (LMA), leaf carboxylation capacity (Vc,max), leaf water content, the degree of isohydry, and predawn Ψleaf. We first investigated how the choice of four commonly used leaf‐level gs models with and without the inclusion of Ψleaf as an additional predictor variable influence the ability to predict gs, and then explored the abiotic (i.e. month, site‐month interaction) and biotic (i.e. tree‐species‐specific characteristics) drivers of slope parameter variation. Our results show that the inclusion of Ψleaf did not improve model performance and that the models that represent the response of gs to vapor pressure deficit performed better than corresponding models that respond to relative humidity. Within each gs model, we found large variation in the slope parameter, and this variation was attributable to the biotic driver, rather than abiotic drivers. We further investigated potential relationships between the slope parameter and the six available plant traits mentioned above, and found that only one trait, LMA, had a significant correlation with the slope parameter (R2 = 0.66, n = 15), highlighting a potential path towards improved model parameterization. This study advances understanding of gs dynamics over seasonal drought, and identifies a practical, trait‐based approach to improve modeling of carbon and water exchange in tropical forests.  相似文献   

19.
The stomatal conductance of several anisohydric plant species, including field-grown sunflower, frequently correlates with leaf water potential (φ1), suggesting that chemical messages travelling from roots to shoots may not play an important role in stomatal control. We have performed a series of experiments in which evaporative demand, soil water status and ABA origin (endogenous or artificial) were varied in order to analyse stomatal control. Sunflower plants were subjected to a range of soil water potentials under contrasting air vapour pressure deficits (VPD, from 0.5 to 2.5 kPa) in the field, in the glasshouse or in a humid chamber. Sunflower plants were also fed through the xylem with varying concentrations of artificial ABA, in the glasshouse and in the field. Finally, detached leaves were fed directly with varying concentrations of ABA under three contrasting VPDs. A unique relationship between stomatal conductance (gs) and the concentration of ABA in the xylem sap (xylem [ABA]) was observed in all cases. In contrast, the relationship between φ1 and gs varied substantially among experiments. Its slope was positive for droughted plants and negative for ABA-fed whole plants or detached leaves, and also varied appreciably with air VPD. All observed relationships could be modelled on the basis of the assumption that φ1 had no controlling effect on gs. We conclude that stomatal control depended only on the concentration of ABA in the xylem sap, and that φ1 was controlled by water flux through the plant (itself controlled by stomatal conductance). The possibility is also raised that differences in stomatal ‘strategy’ between isohydric plants (such as maize, where daytime φ1 does not vary appreciably with soil water status) and anisohydric plants (such as sunflower) may be accounted for by the degree of influence of φ1 on stomatal control, for a given level of xylem [ABA]. We propose that statistical relationships between φ1 and gs are only observed when φ1 has no controlling action on stomatal behaviour.  相似文献   

20.
Studies on the temperature (T) responses of photosynthesis and leaf hydraulic conductance (Kleaf) are important to plant gas exchange. In this study, the temperature responses of photosynthesis and Kleaf were studied in Shanyou 63 (Oryza sativa) and Yannong 19 (Triticum aestivum). Leaf water potential (Ψleaf) was insensitive to T in Shanyou 63, while it significantly decreased with T in Yannong 19. The differential ΨleafT relationship partially accounted for the differing gmT relationships, where gm was less sensitive to T in Yannong 19 than in Shanyou 63. With different gmT and ΨleafT relationships, the temperature responses of photosynthetic limitations were surprisingly similar between the two lines, and the photosynthetic rate was highly correlated with gm. With the increasing T, Kleaf increased in Shanyou 63 while it decreased in Yannong 19. The different KleafT relationships were related to different ΨleafT relationships. When excluding the effects of water viscosity and Ψleaf, Kleaf was insensitive to T in both lines. gm and Kleaf were generally not coordinated across different temperatures. This study highlights the importance of Ψleaf on leaf carbon and water exchanges, and the mechanisms for the gmT and KleafT relationships were discussed.  相似文献   

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