• <tr id="yyy80"></tr>
  • <sup id="yyy80"></sup>
  • <tfoot id="yyy80"><noscript id="yyy80"></noscript></tfoot>
  • 99热精品在线国产_美女午夜性视频免费_国产精品国产高清国产av_av欧美777_自拍偷自拍亚洲精品老妇_亚洲熟女精品中文字幕_www日本黄色视频网_国产精品野战在线观看 ?

    From leaf to whole-plant water use efficiency(WUE) in complex canopies:Limitations of leaf WUE as a selection target

    2015-11-24 12:23:56HiplitoMedrnoMgdlenTomSestiMrtorellJumeFlexsEstherHernndezJonRossellAliciPouJosMrinoEsclonJosefinBot
    The Crop Journal 2015年3期

    Hipólito Medrno,Mgdlen Tomás,Sestià Mrtorell,Jume Flexs, Esther Hernández,Jon Rosselló,Alici Pou,José-Mrino Esclon,Josefin Bot,*

    aResearch Group on Plant Biology under Mediterranean Conditions,Department of Biology,IMEDEA(CSIC-Universitat de les Illes Balears), Carretera de Valldemossa,km 7.5,07122 Palma de Mallorca,Spain

    bInstitute of Life Science,Catholic University of Louvain,Place de l'Université 1,B-1348 Louvain-la-Neuve,Belgium

    From leaf to whole-plant water use efficiency(WUE) in complex canopies:Limitations of leaf WUE as a selection target

    Hipólito Medranoa,1,Magdalena Tomása,Sebastià Martorella,Jaume Flexasa, Esther Hernándeza,Joan Rossellóa(chǎn),Alicia Poub,José-Mariano Escalonaa,Josefina Botaa,*,1

    aResearch Group on Plant Biology under Mediterranean Conditions,Department of Biology,IMEDEA(CSIC-Universitat de les Illes Balears), Carretera de Valldemossa,km 7.5,07122 Palma de Mallorca,Spain

    bInstitute of Life Science,Catholic University of Louvain,Place de l'Université 1,B-1348 Louvain-la-Neuve,Belgium

    A R T I C L E I N F O

    Article history:

    Received 30 December 2014

    Received in revised form 2 April 2015

    Accepted 4 May 2015

    Available online 12 May 2015

    Water use

    Drought

    Intrinsic water use efficiency 13C

    Instantaneous water use efficiency

    Whole plant water use efficiency

    Plant water use efficiency(WUE)is becoming a key issue in semiarid areas,where crop production relies on the use of large volumes of water.Improving WUE is necessary for securing environmental sustainability of food production in these areas.Given that climate change predictions include increases in temperature and drought in semiarid regions, improving crop WUE is mandatory for global food production.WUE is commonly measured at the leaf level,because portable equipment for measuring leaf gas exchange rates facilitates the simultaneous measurement of photosynthesis and transpiration.However, when those measurements are compared with daily integrals or whole-plant estimates of WUE,the two sometimes do not agree.Scaling up from single-leaf to whole-plant WUE was tested in grapevines in different experiments by comparison of daily integrals of instantaneous water use efficiency[ratio between CO2assimilation(AN)and transpiration (E);AN/E]with midday AN/E measurements,showing a low correlation,being worse with increasing water stress.We sought to evaluate the importance of spatial and temporal variation in carbon and water balances at the leaf and plant levels.The leaf position (governing average light interception)in the canopy showed a marked effect on instantaneous and daily integrals of leaf WUE.Night transpiration and respiration rates were also evaluated,as well as respiration contributions to total carbon balance.Two main components were identified as filling the gap between leaf and whole plant WUE:the large effect of leaf position on daily carbon gain and water loss and the large flux of carbon losses by dark respiration.These results show that WUE evaluation among genotypes or treatments needs to be revised.

    ?2015 Crop Science Society of China and Institute of Crop Science,CAAS.Production and hosting by Elsevier B.V.All rights reserved.This is an open access article under the CC BY-NC-ND license(http://creativecommons.org/licenses/by-nc-nd/4.0/).

    1.Introduction

    Water use efficiency(WUE)is an important subject in agriculture in semiarid regions,because of the increasing areas under irrigation and the high water requirements of crops(which consume around 70%of water available to humans).The scarcity of water resources is leading to increasing controversy about the use of water resources by agriculture and industry,for direct human consumption,and for other purposes.Such controversy could be alleviated by increasing crop water use efficiency,so that improving WUE of crops is becoming a main goal for agriculture and food security goals[1–5].Moreover,climate change predictions show clear increases in temperatures(and concomitant increase in potential evapotranspiration)and more frequent episodes of climatic anomalies,such as droughts and heat waves[6,7].All of these climate change phenomena are prevalent in most semiarid areas[8].Consequently,the optimization of water use for crops by improvement of WUE is a challenge for securing agricultural sustainability in semiarid areas.In response to this challenge,a large volume of applied and fundamental research has been focused on optimization of crop water use.

    The water issue is crucial for environmental sustainability of viticulture,because 60% of vineyards are located in semiarid areas and regular water applications are necessary to complete the growth cycle of grapevines.Grapes growth and mature during the driest months,making irrigation scheduling and timing critical[9–11].Consequently,scientific interest in research on grapevine WUE has focused on the evaluation of new irrigation techniques[12–15]and on genetic variation in WUE in grapevine rootstocks or cultivars[16–18] and reflect the social interest and necessity of optimizing water use in viticulture.Fortunately,in most winegrowing regions,the main concern for farmers is not high grape yield but quality.Fruit quality is negatively correlated with high yield[19,20],so that it can be said that high quality yield is generally achieved under suboptimal crop conditions.For this reason,water stress has become a management target to secure high fruit quality,increasing the sustainability of water use by favoring crop quality over quantity.

    WUE can be measured at different scales,ranging from instantaneous measurements on the leaf to more integrative ones at the plant and crop levels(Fig.1).The pros and cons of those different ways to estimate WUE have been discussed elsewhere[21,22],and the decision on the most appropriate way depends on the capacity,facilities,and scale of the specific study. Most studies of WUE are performed on the basis of instantaneous measurements of leaf photosynthesis and transpiration, on the assumption that they are representative of whole-plant WUE,although only a few reports have evaluated WUE at the whole-plantlevel[18,23–25].Comparison between instantaneous and whole-plant values sometimes reveals a clear relationship [10],but often does not.This lack of correspondence is an important limitation to the applicability of the research conducted in this field.Its causes need to be clarified for scaling from single to whole-plant estimates of WUE.

    In the present work we analyze data from multiple experiments identifying sources of environmentally induced leaf WUE variations,showing the importance of both the light environment and dark respiration,often neglected,to whole-plant carbon balance and in turn to whole-plant WUE.

    2.Materials and methods

    2.1.Plant material and treatments

    2.1.1.Field-grown plants

    Fig.1-Different complexity levels for water use efficiency measurements.From leaf to crop level,as from instantaneous to growth-season measurements,there is a progressive integration of different crop production processes and water expenses with different measurement techniques.The double arrows indicate the difficulties in scaling up from leaf to plant level.

    A field experiment was conducted in the experimental field of the University of Balearic Islands(Majorca,Spain)on grapevines of the cultivars Tempranillo and Grenache duringsummer 2012,as described by Martorell et al.[26].Plants were five years old(planted in 2007)grafted onto 110-Richter rootstock and planted 1 m apart in rows 2.5 m apart.They were trained in a bilateral double cordon having between 10 and 12 canes per plant in 2012.Two irrigation treatments were applied:well watered(WW),consisting of approximately 9 L day?1plant?1, and water stress(WS),consisting of withholding irrigation for the entire summer.Predawn leaf water potential(Ψpd)was used as a stress indicator.Ψpdwas measured monthly(June,July and August)with a Scholander pressure chamber(Soil Moisture Equipment Corp.,Santa Barbara,CA).Four replicates per treatment and cultivar were measured.The WW treatment maintained Ψpdbetween?0.16 and?0.27 MPa in Grenache and between?0.16 and?0.30 MPa in Tempranillo.The WS treatment reduced Ψpdin August to a minimum of?0.85 MPa in Grenache and?0.53 MPa in Tempranillo.

    2.1.2.Potted plants

    Seven different cultivars of grapevine subjected to well-watered and water-stressed conditions were studied in three different experiments performed in three consecutive years(summer 2008,2009,and 2010)at the University of Balearic Islands (Majorca,Spain),as described in Tomás et al.[18].Briefly, ungrafted plants were grown outdoors in 15-L pots in a mixture of organic substrate and perlite(3:1).The cultivars Malvasia of Banyalbufar,Cabernet Sauvignon,Grenache,and Tempranillo were studied in all three years.Escursac,Manto Negro,and Pinot Noir were studied only in 2010.

    Environmental conditions were recorded during the experiment using a meteorological station(Meteodata 3000,Geónica SA,Madrid,Spain).In general,the climatic variables were very similar in the three experimental years with small differences in mean air temperature,which ranged from 25.7°C in 2008 to 24.2°C in 2010.Total daily potential evapotranspiration was not significantly different among the three experimental periods (5.3–5.6 L m?2day?1).Well-watered plants were irrigated to field capacity throughout the experiment.

    Moderate water stress level was sustained for three weeks to maintain leaf maximum daily stomatal conductance values (gs),around 0.05 mol CO2m?2s?1.Once the desired level of water stress was reached,plants were maintained under constant water stress for three weeks by daily replacement of the exact amount of water consumed,determined by pot weight.The imposition of water stress treatment produced large reductions in soil water content(SWC calculated as follows:(pot weight?minimum pot weight)/(maximum pot weight?minimum pot weight)×100),from 70–90%(well irrigated)down to 13–22%.

    An additional experiment was performed in September of 2010 in the same location,as described in Escalona et al.[27]. Ten-year-old grapevine plants(cv.Tempranillo)grafted onto 110-Richter rootstock,were grown outside in 60-L containers in a mix of sand,Prohumin(Projar SA,Valencia,Spain) horticultural substrate,and perlite(1:1:1).The surfaces of containers were covered with a thin layer of perlite and sealed with plastic film(held with a rubber band around the edge of each container)to minimize water losses by direct evaporation.Two treatments were imposed:(i)five plants were maintained at field capacity throughout the experiment by daily irrigation and(ii)five plants were subjected to progressive drought stress by withholding irrigation.Stem water potential (Ψstem)was used as a stress indicator.Leaves were sealed in a plastic bagand covered with aluminumfoil.After 1 h,Ψstemwas measured using a Scholander pressure chamber(see above). Different levels of water stress were obtained with time; moderate drought stress was achieved by day 4,when plants showed Ψstemvalues of?0.8 MPa,and severe stress by day 7, when Ψstemreached?1.34 MPa.The plants were maintained outside during the growing season.At the outset of the experiment,plants showed 8 shoots of 1.5 m length with about 20 leaves per shoot.

    2.2.Gas exchange measurements

    Instantaneous gas exchange measurements were made on four to six recently fully expanded leaves in the upper part of the canopy for each variety and treatment between 10:00 and 12:00 h using an open gas exchange system(LI-6400;LI-COR, Inc.,Lincoln,Nebraska USA).

    Measurements of net CO2assimilation(AN),gs,and transpiration(E)were performed at saturating red light(1500 μmol m?2s?1) achieved with the red LED lamp of the system,with an additional 10%of blue light to maximize stomatal opening, and 400 μmol CO2mol?1in the cuvette.Air temperature and humidity in the chamber was set to match environmental conditions,in consequence of which leaf temperature ranged between 28 and 34°C depending on leaf water status.

    Gas exchange measurements were made in leaves located at 14 different positions in the canopy(lower,medium and upper parts of east and west sides and internal leaves)on August 23, 2012 in five-year-old Tempranillo plants(in the field experiment) using the same(LI-6400)open-flow gas exchange system equipped with a clear chamber(LI-6400-08).Air temperature and humidity in the chamber was set to match ambient and CO2concentration was set at 400 mol mol?1.

    Intrinsic water use efficiency(AN/gs)was calculated as the ratio between ANand gs,and instantaneous water use efficiency (AN/E)between ANand E.

    2.3.Night transpiration and respiration rates

    Night transpiration was measured on Tempranillo potted plants as described in Escalona et al.[27].Briefly,gas exchange measurements were performed using the LI-6400 instrument equipped with a 6 cm2chamber.Measurements were performed at 400 mol CO2mol?1of air and at low airflow rates(150 mol air s?1)on three leaves per plant(15 replicates per treatment)every 2 h during the entire nighttime period, starting 1 h after sunset(19:30 solar time)and finishing 1 h before dawn(05:30 solar time).

    Respiration rates of plant organs and plant carbon balance estimation were performed in potted plants of Tempranillo and Grenache cultivars during summer 2010 as described in Escalona et al.[28].

    2.4.Whole plant water use efficiency and carbon isotope composition

    In potted plants of seven cultivars(see Plant material and treatments),four plants per cultivar were harvested todetermine initial whole-plant biomass.Similarly,four plants per cultivar and treatment were harvested at the end of the experiment.Leaves,shoots and roots per plant were separated and dried in an oven at 60°C to obtain dry weight.The total biomass increase during the experiment was estimated as the difference between the whole-plant dry weights at the beginning and end of the experiment.

    Plant water consumed over the three-week period was estimated from the sum of the daily water consumption as previously described.

    Whole plant WUE was determined as follows:

    For carbon isotope composition,six young leaves per cultivar and treatment from different plants,developed after the outset of the stress treatment,were sampled at the end of the experiment.They were dried for 48 h at 60°C and ground into powder.Subsamples of 2 mg were analyzed for isotope ratio(δ13C)as a long-term indicator of WUE.The samples were combusted in an elemental analyzer(Carlo-Erba,Rodano, Italy),CO2was separated by chromatography and directly injected into a continuous-flow isotope ratio mass spectrometer(Thermo Finnigan Delta Plus,Bremen,Germany).Peach leaf standards(NIST 1547)were run every eight samples.δ13C was calculated as follows:

    δ13C values were referenced to a Pee Dee Belemnite standard.

    3.Results and discussion

    3.1.Variation in WUE over time

    A literature survey of plant WUE shows that WUE determinations rely on direct measurement of instantaneous gas exchange rates(photosynthesis and transpiration)at the leaf level with portable equipment.Usually,such measurements are taken on recently fully expanded leaves,well light-exposed,around midmorning because in most cases this time yields the highest values for AN,gs,and E.However,as Fig.2 shows, there is large variation in“intrinsic”water use efficiency (estimated as AN/gs)throughout the day,as measured under field conditions.The figure shows that at the typical measurement time(midmorning)AN/gsvalues range from 50 to 70 μmol CO2mol?1H2O,but that afterwards AN/gsvalues were higher or lower and that these daily changes are even higher under water stress.The evidence of these daily time changes calls into question the widely accepted principle of optimization of resources by the plant,showing how daily variations in environmental and leaf conditions correspond to large changes in physiological parameters.The extent to which the typically measured values are representative of the whole dayAN/gsisnotunder discussion,although obviously integration over the full day would more accurately represent the actual leaf WUE.Measurement limitations always influence the decision between greater numbers of more comparable measurements and more accurate but restrictive measurements.The limitation imposed by daily variation in WUE has been shown by Medrano et al.[29]with plots of midmorning values of AN/gsagainst whole-day integrals(asμmol CO2mol?1H2O day?1m?2)for different grapevine genotypes.The correspondences were high or low,depending on the experiment.

    Fig.2-Diurnal time variation in intrinsic WUE(as AN/gs). Diurnal time cycles of grapevine(cultivar Tempranillo)under irrigation(filled symbols)and moderate water-stress conditions(open symbols).Plants were grown outdoors in the field during summer in Mallorca(Balearic Islands,Spain). Values are averages of 5 replicates±SE.

    Along with diurnal time effects,there is seasonal variation in leaf WUE as a consequence of both changing environmental conditions and the physiological changes expected with leaf aging,which modifies leaf photosynthesis and transpiration.Fig.3 shows how these changes in grapevine leaves modify intrinsic WUE from early growth to harvest.Under irrigation,the midmorning values of AN/gs,measured in field-grown Grenache and Tempranillo plants,changed from 40 to 80 μmol CO2mol?1H2O,similar in the two varieties. However,the increase in AN/gsin response to moderate water stress is greater for Grenache than for Tempranillo from veraison to ripening and harvest time,and the reputation of Grenache as a more drought-resistant variety is more clearly corroborated in the latter growth periods.

    3.2.Spatial variation in leaf WUE in complex canopies:the case of grapevine

    Fig.3-Variation of intrinsic WUE(as AN/gs)with phenology. Intrinsic WUE variation throughout the growing period in two different grapevine cultivars grown in the field from bloom to harvest period.Black symbols,well-watered plants; white symbols,water-stressed plants.Cultivars are Tempranillo(squares)and Grenache(triangles).Values are averages of 5 replicates±SE.

    In complex canopies,the light intercepted by an individual leaf is highly dependent on the leaf position and the canopy geometry.In the grapevine,the trellis system and row orientation provide differential light exposure for different leaf positions in the canopy,corresponding to differences in microclimate that clearly affect the daily time course of leaf gas exchange rates.The effect of leaf position on integral daily carbon gain was reported by Escalona et al.[30],showing large variation from top layers of the canopy receiving 100%of incoming light to lower positions receiving only 25%,and showing that inner shaded leaves received only around 5%of incoming light.This differential light and microclimate environment caused large changes in the daily time courses of photosynthesis and transpiration but also in daily and seasonal integrals,leading to large variation in carbon gain and water consumption among different positions of the canopy. Concerning leaf WUE,reanalyzing these data,Medrano et al.[29], showed that both instantaneous and daily integrals of leaf WUE (as integrals of AN/gsor AN/E values)were also highly dependent on the microclimate environment of each leaf position and that WUE values of upper locations were double those of lower ones. These variations were similar or even higher under moderate and severe water stress.In fact,daily leaf WUE proved to be highly determined by the daily intercepted light at each leaf position(with a R2of 0.98 for irrigated plants).Fig.4 shows the effect of leaf position on WUE at multiple positions in the canopy (14).The average values of typical midmorning measurements WUE(as AN/gs)for well-irrigated vines showed a similar tendency,with clear differentiation(expressed as values three to four times higher)between the east and west sides of the canopy.

    Those results show large spatial variation of WUE in the canopy as well as the importance of this complexity for the evaluation of plant WUE on the basis of instantaneous measurements of typical fully exposed leaves in specific locations.These results also provide an interesting example of the fine responses of leaves to environmental variation, showing that leaf capacity to regulate photosynthesis and transpiration results in large variation in WUE,calling into question the leaf gas-exchange rate optimization theory[31]. Although comparative studies on the basis of WUE measurements in a single leaf are useful and provide an affordable way to compare genotypes and agronomic practices,the relationships among these standard values and whole-plant values are not simple,because of the complexity of the canopy and the differential responses of the leaf to cumulative daily irradiance.

    Fig.4-Variation in intrinsic WUE(as AN/gs)with leaf position in the canopy.Intrinsic WUE(AN/gs,μmol CO2mol?1H2O) measured at 14 positions throughout the canopy at midday in five-year-old grapevines of cv.Tempranillo under field conditions.Values are means of four replicates±SE measured in August 2012.

    3.3.From leaf to whole plant WUE:effects of night transpiration and respiration rate

    Fig.5-Night transpiration rates in irrigated and water-stressed grapevines.Leaf night transpiration rates during a typical dark period in 10-year-old potted Tempranillo grapevines grown under irrigation(black symbols)and water stress(white symbols).

    Besides the difficulties of extrapolating to whole-plant photosynthesis and transpiration from instantaneous single-leaf measurement,whole-plant WUE measured as biomass increase with water used is strongly dependent on other physiological processes determining WUE:respiration losses and night transpiration.

    Night transpiration has recently been reviewed and measured,proving to be non-negligible and possibly markedly reduced under water stress.Also,under certain circumstances nighttime transpiration can account for 10%of daily transpiration losses[27,32].Fig.5 shows,as an example,nighttime leaf transpiration rates of irrigated and water-stressed 10-year-old potted Tempranillo plants,showing rates of around 10%of daily transpiration(data not shown)and threefold higher in irrigated than in water-stressed grapevines.As already reported by our group[27],these differences between treatments cannot be explained by epicuticular changes.Using whole-plant mini-lysimeters,we also showed that on very humid nights these losses are nearly compensated by dew income[27].In any case,these water losses reduce the expected daily WUE of the whole plant.

    Fig.6-Carbon balance as affected by respiration components.Contribution as percentage of total carbon gain of several respiratory components of grapevines(cultivars Grenache and Tempranillo).Net carbon gain is expressed as the remaining percentage.

    Fig.7-Relationships between water use efficiency(WUE)measured at leaf and whole-plant levels.Relationships between intrinsic WUE(AN/gs)and whole-plant WUE(WUEWP)in 2008(A),2009(B),and 2010(C).Relationships between instantaneous WUE(AN/E)and WUEWPmeasured in 2008(D),2009(E),and 2010(F).Relationship between leaf δ13C and WUEWPfrom values obtained in three different experiments,2008(G),2009(H),and 2010(I).AN/gsand AN/E were measured in midmorning during the experimental periods and leaf δ13C was measured at the end of the experimental periods.Black symbols,control plants; white symbols,water-stressed plants.Cultivars are represented as follows:Tempranillo(square),Malvasia of Banyalbufar (circle),Grenache(inverted triangle),Cabernet Sauvignon(upright triangle),Callet(diamond),Richter-110(cross),Escursac (plus),Manto Negro(hexagon),and Pinot Noir(star).Values are means±SE of six replicates.WUEWPwas measured at the end of each experiment.Values are averages of 4 replicates±SE.

    With respect to carbon losses by respiratory processes, there is complexity derived from respiration rate variation with environmental conditions and plant growth and development.A main limitation to evaluating the effect on carbon balance of variation in respiration rate is the paucity of studies on dark respiration.Estimates of plant respiration are often obtained by measurement in organs such as leaves,shoots,and roots,but most reports have focused on leaf respiration.However,it is well known that the largest respiratory losses come from the root system,presenting great difficulty for accurate determination under field conditions.Still,for grapevines there are some reports on root respiration rate[28,33–35]based on calculation for pot-and field-grown plants and assuming specific soil respiration activities for the latter.Similarly to other parameters, leaf respiration rates showed large variation with canopy position(for leaves),age(for shoots and fruits),and plant water status[28].The relative magnitudes of respiration losses are shown in Fig.6,showing that,in Tempranillo and Grenache potted plants,respiration losses represent around 33%of total carbon gain for irrigated plants and around 45%for water-stressed plants.Among those respiration losses,there is clear variation linked to water status and also differences between the two sampled varieties.In general,root respiration losses seem to be the main losses,followed closely by fruit and leaf respiration losses.

    Overall,these results confirmed the importance of respiratory losses for understanding plant carbon balance,but also for better understanding dark respiration as the largest unknown factor relating leaf instantaneous and whole-plant water use efficiency.Certainly,as Fig.6 shows,respiration rates are not constant but show wide variation with water status and variety.

    3.4.The missing key:identifying a more representative indicator of WUE

    As described above,there are different sources of variation in carbon gain and water loss,from the single leaf to the whole-plant,which can affect the correspondence betweenleaf and whole plant determinations of WUE.We have shown the large variation of photosynthesis and transpiration with leaf diurnal time courses and seasonal variation,but there is also marked variation with leaf position and a large effect of respiratory losses.As shown by Tomás et al.[36],the relative importance of this canopy complexity and plant respiration of grapevines can be weighted on the basis of collected data for irrigated Tempranillo vines.For this analysis the effect of canopy complexity[29,30]was assessed,considering the potential maximum values of AN,E,and whole plant WUE as those that would be achieved by a plant if all of its leaves were fully exposed to the sun throughout the day.These theoretical carbon gains(AN)and water losses(E)were defined as 100%,so that the theoretical maximum AN/E ratio is 1.Taking into account the canopy location effect,the potential carbon gain by unit of leaf area falls to around 47%.With respect to transpiration,the canopy effect reduces water losses by around 36%.Consequently,the plant WUE would be expected to be around 0.75 of the“potential”whole-plant WUE.Introducing the effect of plant respiration,the carbon losses from respiration in roots,fruits,leaves,and shoots represent from 30%to 45%of the carbon fixed.Finally,considering the night transpiration component[27],transpiration increases,so that recalculating net carbon gain and transpiration losses yields a final WUE of the plant around 33%of the theoretically estimated WUE.Fig.7 shows the general lack of relationship between leaf-level estimates of WUE(AN/gs,AN/E,or δ13C),with whole-plant biomass-based WUE(WUEWP),using data from seven grapevine cultivars over three consecutive years and two water-availability conditions.Among all of these studied combinations,only a few showed a clear correspondence between single-leaf and whole-plant measurements(Fig.7A,C, G).For instance,in two of the three experimental periods,AN/gswas significantly and positively correlated with WUEWPin non-irrigated plants,but the correlation was lost when irrigated plants were also considered.For δ13C,a single significant negative correlation with WUEWPwas observed when all data in the first of the three experiments were pooled,but no such correlation was observed in the other two experiments. Moreover,while water stress results in increased leaf-level WUE in all cases,its effects on WUEWPare variable,from decreases in most cultivars in 2008 and some in 2010,to no changes or increases,depending on the cultivar,in 2009 and 2010.Genetic variability in WUE at different levels was recently reviewed[36,37]and the predicted causes of discrepancies between WUE values at leaf level and WUEWP[10,29]were associated with all of the components analyzed in the present study:the complexity of light interception,night transpiration, and respiratory losses.Thus,these are major limitations to choosing a single selection criterion to rate the WUE of a given genotype.This difficulty is a serious handicap to conducting selection programs for this character.

    4.Conclusion

    The reported lack of correspondence among leaf gas exchange parameters and whole plant carbon and water balances imposes a severe limitation on the accurate measurement of treatment and/or genotype effects on whole-plant WUE under field conditions.It is thus necessary to fill the gaps in scaling from single-leaf to whole-plant estimates of WUE to better understand the underlying processes leading to the variety of responses.This variety illustrates the diversity of single-leaf vs.whole-plant WUE relationships.The reported data and discussion clearly show that a more intensive and extensive research effort is needed to improve the representativeness of typical sampling procedures in evaluating whole-plant WUE.

    Acknowledgments

    This work was performed with financial support from the Spanish Ministry of Science and Technology(project AGL2011-30408-C04-01)and from Conselleria de Educación, Cultura y Universidades(Govern de les Illes Balears)and the European Social Fund through the ESF Operational Programme for the Balearic Islands 2013–2017(project PD/027/2013).

    [1]J.L.Araus,The problems of sustainable water use in the Mediterranean and research requirements for agriculture, Ann.Appl.Biol.144(2004)259–272.

    [2]X.P.Deng,L.Shan,H.Zhang,N.C.Turner,Improving agricultural water use efficiency in arid and semiarid areas of China,Agric.Water Manag.80(2006)23–40.

    [3]S.Geerts,N.Raes,Deficit irrigation as an on-farm strategy to maximize crop water productivity in dry areas,Agric.Water Manag.96(2009)1275–1284.

    [4]N.Katerji,M.Mastrorilli,G.Ranab,Water use efficiency of crops cultivated in the Mediterranean region:review and analysis,Eur.J.Agron.28(2008)493–507.

    [5]J.I.L.Morison,N.R.Baker,P.M.Mullineaux,W.J.Davies, Improving water use in crop production,Philos.T.R.Soc,B 363(2008)639–658.

    [6]IPCC,T.F.Stocker,Q.Dahe,G.K.Plattner,M.Tignor,S.K.Allen, J.Boschung,A.Nauels,Y.Xia,V.Bex,P.M.Midgley,Climate change 2013:the physical science basis,Working Group I Contribution to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change,Cambridge University Press,Cambridge and New York,2013.

    [7]H.G.Jones,R.A.Vaughan,Remote Sensing of Vegetation: Principles,Techniques,and Applications,Oxford University Press,Oxford,2010.

    [8]S.M.Vicente-Serrano,J.I.Lopez-Moreno,S.Beguería,J. Lorenzo-Lacruz,A.Sanchez-Lorenzo,J.M.García-Ruiz,C. Azorin-Molina,E.Morán-Tejeda,J.Revuelto,R.Trigo,F. Coelho,F.Espejo,Evidence of increasing drought severity caused by temperature rise in Southern Europe,Environ.Res. Lett.9(2014)044001.

    [9]M.M.Chaves,T.P.Santos,C.R.Souza,M.F.Ortu?o,M.L. Rodrigues,C.M.Lopes,J.P.Maroco,J.S.Pereira,Deficit irrigation in grapevine improves water-use-efficiency without controlling vigour and production quality,Ann.Appl. Biol.150(2007)237–252.

    [10]J.Flexas,J.Galmés,A.Gallé,J.Gulías,A.Pou,M.Ribas-Carbó, M.Tomás,H.Medrano,Improving water use efficiency in grapevines:potential physiological targets for biotechnological improvement,Aust.J.Grape Wine Res.161 (2010)106–121.

    [11]L.E.Williams,J.E.Ayars,Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy,Agric.For.Meteorol.132(2005)201–211.

    [12]V.O.Sadras,Does partial root-zone drying improve irrigation water productivity in the field?A meta-analysis,Irrig.Sci.27 (2009)183–190.

    [13]L.E.Williams,Interaction of rootstock and applied water amounts at various fractions of estimated evapotranspiration (ETc)on productivity of Cabernet sauvignon,Aust.J.Grape Wine Res.3(2010)434–444.

    [14]M.M.Chaves,O.Zarrouk,R.Francisco,J.M.Costa,T.Santos, A.P.Regalado,L.Rodrigues,C.M.Lopes,Grapevine under deficit irrigation:hints from physiological and molecular data,Ann.Bot.105(2010)661–676.

    [15]P.Romero,J.I.Fernández-Fernández,A.Martinez-Cutillas, Physiological thresholds for efficient regulated deficitirrigation management in wine grapes grown under semiarid conditions,Am.J.Enol.Vitic.61(2010)300–312.

    [16]J.Satisha,G.S.Prakash,R.Venugopalan,Statistical modeling of the effect of physio-biochemical parameters on water use efficiency of grape varieties,rootstocks and their stionic combinations under moisture stress conditions,Turk.J. Agric.For.30(2006)261–271.

    [17]M.M.Alsina,F.de Herralde,X.Aranda,R.Save,C.Biel,Water relations and vulnerability to embolism are not related: experiments with eight grapevine cultivars,Vitis 46(2007) 1–6.

    [18]M.Tomás,M.Medrano,A.Pou,J.M.Escalona,S.Martorell,M. Ribas-Carbó,J.Flexas,Water use efficiency in grapevine cultivars:effects of water stress at leaf and whole plant level, Aust.J.Grape Wine Res.18(2012)164–172.

    [19]P.Romero,R.Mu?oz,F.Del Amor,E.Valdes,J.I.Fernández,A. Martinez-Cutillas,Regulated deficit irrigation based upon optimum water status improves phenolic composition in Monastrell grapes in wines,Agric.Water Manag.121(2013) 85–101.

    [20]L.E.Williams,M.A.Matthews,Irrigation of agricultural crops. Agronomy monographs no.30,in:B.J.Stewart,D.R.Nielsen (Eds.),Experimental Agriculture,Cambridge University Press, Cambridge 1990,pp.1019–1055.

    [21]H.Medrano,J.Gulías,M.Chaves,J.Galmés,J.Flexas, Photosynthesis water-use efficiency,in:J.Flexas,F.Loreto,H. Medrano(Eds.),Terrestrial Photosynthesis in a Changing Environment,A Molecular,Physiological and Ecological ApproachCambridge University Press,Cambridge 2012, pp.529–543.

    [22]H.Medrano,J.Flexas,M.Ribas-Carbó,J.Gulías,Measuring water use efficiency in grapevines,in:S.Delrot,H.Medrano, E.Or,L.Bavaresco,S.Grando(Eds.),Methodologies and Results Grapevine Research,Springer,Germany 2010, pp.57–60.

    [23]M.R.Gibberd,R.R.Walker,D.H.Blackmore,A.G.Condon, Transpiration efficiency and carbon-isotope discrimination of grapevines grown under well-watered conditions in either glasshouse or vineyard,Aust.J.Grape Wine Res.7(2001) 110–117.

    [24]S.Poni,F.Bernizzoni,S.Civardi,M.Gatti,D.Porro,F.Camin, Performance and water-use efficiency(single-leaf vs.wholecanopy)of well-watered and half-stressed split-root Lambrusco grapevines grown in Po Valley(Italy),Agric. Ecosyst.Environ.129(2009)97–106.

    [25]J.M.Tarara,J.E.Pérez-Pe?a,R.P.Schreiner,M.Keller,P. Smithyman,Net carbon exchange in grapevine canopies responds rapidly to timing and extent of regulated deficit irrigation,Funct.Plant Biol.38(2011)386–400.

    [26]S.Martorell,H.Medrano,M.Tomás,J.M.Escalona,J.Flexas,A. Díaz-Espejo,Plasticity of vulnerability to leaf hydraulic dysfunction during acclimation to drought in grapevines:an osmotic-mediated process,Physiol.Plant.153(2015)381–391.

    [27]J.M.Escalona,S.Fuentes,M.Tomás,S.Martorell,J.Flexas,H. Medrano,Responses of leaf night transpiration to drought stress in Vitis vinifera L,Agric.Water Manag.118(2013)50–58.

    [28]J.M.Escalona,M.Tomás,S.Martorell,H.Medrano,M. Ribas-Carbó,J.Flexas,Carbon balance in grapevines under different soil water supply:importance of whole plant respiration,Aust.J.Grape Wine Res.18(2012)308–318.

    [29]H.Medrano,A.Pou,M.Tomás,S.Martorell,J.Gulias,J.Flexas, J.M.Escalona,Average daily light interception determines leaf water use efficiency among different canopy locations in grapevine,Agric.Water Manag.114(2012)4–10.

    [30]J.M.Escalona,J.Flexas,J.Bota,H.Medrano,Distribution of leaf photosynthesis and transpiration within grapevine canopies under different drought conditions,Vitis 42(2003) 57–64.

    [31]T.N.Buckley,S.Martorell,A.Díaz-Espejo,M.Tomás,H. Medrano,Is stomatal conductance optimized over both time and space in plant crowns?A field test in grapevine(Vitis vinifera),Plant Cell Environ.27(2014)1–15.

    [32]S.Fuentes,R.De Bei,M.Collins,J.M.Escalona,H.Medrano, S.D.Tyerman,Night time responses to water supply in grapevines(Vitis vinifera L.)under deficit irrigation and partial root-zone drying,Agric.Water Manag.138(2014)1–9.

    [33]K.Morinaga,S.Imai,H.Yakushiji,Y.Koshita,Effects of fruit load on partitioning of15N and13C,respiration,and growth of grapevine roots at different fruit stages,Sci.Hortic.97(2003) 239–253.

    [34]Z.W.Dai,P.Vivin,F.Barrieu,N.Ollat,S.Delrot,Physiological and modelling approaches to understand water and carbon fluxes during grape berry growth and quality development:a review,Aust.J.Grape Wine Res.16(2010)70–85.

    [35]N.Franck,J.P.Morales,D.Arancibia-Avendano,V.G.de Cortazar,J.F.Pérez-Quezada,A.Zurita-Silva,C.Pastenes, Seasonal fluctuations in Vitis vinifera root respiration in the field,New Phytol.192(2011)939–951.

    [36]M.Tomás,H.Medrano,J.M.Escalona,S.Martorell,A.Pou,M. Ribas-Carbó,J.Flexas,Genetic variability in water use efficiency in grapevines,Environ.Exp.Bot.103(2014)148–157.

    [37]H.Medrano,M.Tomás,S.Martorell,J.M.Escalona,A.Pou,S. Fuentes,J.Flexas,J.Bota,Improving water use efficiency of vineyards in semiarid regions:a review,Agron.Sustain.Dev. 35(2015)499–517.

    *Corresponding author.Tel.:+34 971173168.

    E-mail address:j.bota@uib.es(J.Bota).

    Peer review under responsibility of Crop Science Society of China and Institute of Crop Science,CAAS.1Both authors contributed equally to the present paper.

    http://dx.doi.org/10.1016/j.cj.2015.04.002

    2214-5141/?2015 Crop Science Society of China and Institute of Crop Science,CAAS.Production and hosting by Elsevier B.V.All rights reserved.This is an open access article under the CC BY-NC-ND license(http://creativecommons.org/licenses/by-nc-nd/4.0/).

    在线天堂中文资源库| 精品一区二区免费观看| 91久久精品国产一区二区三区| 免费大片18禁| 黄色视频在线播放观看不卡| 男女午夜视频在线观看 | 成人手机av| 精品国产乱码久久久久久小说| 亚洲国产欧美在线一区| 99九九在线精品视频| 下体分泌物呈黄色| 自线自在国产av| 久久精品熟女亚洲av麻豆精品| 岛国毛片在线播放| 伊人亚洲综合成人网| 亚洲精品,欧美精品| 国产精品秋霞免费鲁丝片| 搡老乐熟女国产| 欧美 亚洲 国产 日韩一| 国产精品久久久久久久久免| 国产成人一区二区在线| 欧美亚洲 丝袜 人妻 在线| 黑人高潮一二区| 卡戴珊不雅视频在线播放| 97精品久久久久久久久久精品| 丁香六月天网| 亚洲成人av在线免费| 久久99蜜桃精品久久| 九色成人免费人妻av| 亚洲婷婷狠狠爱综合网| 成年av动漫网址| 欧美老熟妇乱子伦牲交| freevideosex欧美| 一级毛片电影观看| 亚洲人与动物交配视频| 两个人免费观看高清视频| 精品国产露脸久久av麻豆| 黄片无遮挡物在线观看| 少妇被粗大猛烈的视频| 亚洲情色 制服丝袜| 中国国产av一级| av国产久精品久网站免费入址| 99热这里只有是精品在线观看| 国产色婷婷99| 永久免费av网站大全| 精品午夜福利在线看| 最后的刺客免费高清国语| 欧美亚洲日本最大视频资源| 少妇高潮的动态图| 日韩不卡一区二区三区视频在线| 久久韩国三级中文字幕| 高清毛片免费看| 哪个播放器可以免费观看大片| 一级毛片电影观看| 麻豆精品久久久久久蜜桃| 男女无遮挡免费网站观看| 国产av码专区亚洲av| 日韩精品有码人妻一区| 亚洲高清免费不卡视频| 成年av动漫网址| 日本黄色日本黄色录像| 欧美日韩成人在线一区二区| 9色porny在线观看| 蜜桃国产av成人99| 岛国毛片在线播放| 国产极品粉嫩免费观看在线| 亚洲婷婷狠狠爱综合网| 亚洲第一av免费看| 日韩av不卡免费在线播放| 国产一区亚洲一区在线观看| 亚洲国产色片| 天天影视国产精品| 中文天堂在线官网| 亚洲av电影在线观看一区二区三区| 男人爽女人下面视频在线观看| 国产日韩欧美在线精品| 午夜福利网站1000一区二区三区| 成人影院久久| 久久99热6这里只有精品| 亚洲欧美成人精品一区二区| 69精品国产乱码久久久| 国产免费一区二区三区四区乱码| 青春草视频在线免费观看| 大话2 男鬼变身卡| 欧美精品av麻豆av| 国产精品久久久久久久久免| 不卡视频在线观看欧美| 精品国产一区二区久久| 婷婷色综合www| 免费黄频网站在线观看国产| 天堂中文最新版在线下载| 最近最新中文字幕免费大全7| 91精品伊人久久大香线蕉| av线在线观看网站| 免费观看av网站的网址| 黑人猛操日本美女一级片| 亚洲人与动物交配视频| 国产在线视频一区二区| 日本vs欧美在线观看视频| 久久午夜福利片| 精品人妻一区二区三区麻豆| 天天操日日干夜夜撸| 大片电影免费在线观看免费| 日韩 亚洲 欧美在线| 欧美变态另类bdsm刘玥| 久久99蜜桃精品久久| 9色porny在线观看| 国产爽快片一区二区三区| 亚洲国产日韩一区二区| 搡女人真爽免费视频火全软件| 嫩草影院入口| 久久国产精品男人的天堂亚洲 | 热re99久久国产66热| 韩国精品一区二区三区 | 亚洲精品成人av观看孕妇| 国产又色又爽无遮挡免| 69精品国产乱码久久久| 9191精品国产免费久久| 美女中出高潮动态图| 母亲3免费完整高清在线观看 | 日日爽夜夜爽网站| 国产日韩欧美亚洲二区| 日韩av不卡免费在线播放| 国产精品一区二区在线不卡| 久久久久久人妻| 国产精品一区二区在线观看99| 美女国产视频在线观看| 人妻少妇偷人精品九色| 不卡视频在线观看欧美| 亚洲成国产人片在线观看| 99久久精品国产国产毛片| 国产精品麻豆人妻色哟哟久久| 一级毛片黄色毛片免费观看视频| 日韩av免费高清视频| 97人妻天天添夜夜摸| 精品一品国产午夜福利视频| 亚洲天堂av无毛| 久久久久久久精品精品| 乱人伦中国视频| 大陆偷拍与自拍| 最近最新中文字幕免费大全7| 大香蕉97超碰在线| 另类亚洲欧美激情| 日本午夜av视频| 啦啦啦视频在线资源免费观看| 成年av动漫网址| 美女脱内裤让男人舔精品视频| 精品一区二区免费观看| 国产精品一区www在线观看| 久久久精品免费免费高清| 亚洲一区二区三区欧美精品| 嫩草影院入口| 亚洲丝袜综合中文字幕| 制服诱惑二区| 一级毛片我不卡| 午夜影院在线不卡| 亚洲精品日韩在线中文字幕| 久久韩国三级中文字幕| 欧美变态另类bdsm刘玥| 日韩伦理黄色片| videossex国产| 一个人免费看片子| 国产精品偷伦视频观看了| 日日爽夜夜爽网站| 午夜免费观看性视频| av在线观看视频网站免费| 日本av手机在线免费观看| 欧美成人午夜免费资源| 精品熟女少妇av免费看| 欧美国产精品一级二级三级| 久热久热在线精品观看| 国产女主播在线喷水免费视频网站| 亚洲国产精品一区三区| 亚洲国产毛片av蜜桃av| 欧美日韩视频精品一区| 亚洲欧美清纯卡通| 十八禁网站网址无遮挡| 久久久久视频综合| 少妇熟女欧美另类| 亚洲经典国产精华液单| 成人影院久久| 一个人免费看片子| 日本午夜av视频| 日韩免费高清中文字幕av| 人妻 亚洲 视频| 成人黄色视频免费在线看| 亚洲高清免费不卡视频| 熟女av电影| 搡女人真爽免费视频火全软件| 在线 av 中文字幕| 看十八女毛片水多多多| 搡老乐熟女国产| 亚洲精品自拍成人| 日韩三级伦理在线观看| av国产久精品久网站免费入址| 老女人水多毛片| 亚洲精品日韩在线中文字幕| 亚洲综合精品二区| 99国产精品免费福利视频| 日本vs欧美在线观看视频| 最近2019中文字幕mv第一页| 99视频精品全部免费 在线| 校园人妻丝袜中文字幕| 欧美 亚洲 国产 日韩一| 免费看不卡的av| 最黄视频免费看| 99久久精品国产国产毛片| a级片在线免费高清观看视频| 又大又黄又爽视频免费| 亚洲精品第二区| 国产免费现黄频在线看| 成人18禁高潮啪啪吃奶动态图| av线在线观看网站| 欧美精品一区二区免费开放| www日本在线高清视频| 午夜福利,免费看| 亚洲国产精品专区欧美| 一级黄片播放器| 亚洲av电影在线进入| 亚洲欧洲日产国产| 涩涩av久久男人的天堂| 汤姆久久久久久久影院中文字幕| 欧美人与性动交α欧美软件 | 91精品伊人久久大香线蕉| 精品一区二区三卡| 婷婷色综合大香蕉| 一本大道久久a久久精品| 宅男免费午夜| 中文精品一卡2卡3卡4更新| 在线观看www视频免费| 波野结衣二区三区在线| 黄色怎么调成土黄色| 欧美精品高潮呻吟av久久| 只有这里有精品99| 9热在线视频观看99| 欧美日本中文国产一区发布| 国产男人的电影天堂91| 亚洲国产精品国产精品| 2022亚洲国产成人精品| 日韩欧美精品免费久久| 色5月婷婷丁香| 色婷婷av一区二区三区视频| a 毛片基地| 亚洲国产日韩一区二区| av一本久久久久| 男人舔女人的私密视频| 91午夜精品亚洲一区二区三区| 毛片一级片免费看久久久久| 亚洲国产精品999| 美女国产视频在线观看| 亚洲精品456在线播放app| 中文字幕人妻丝袜制服| 黄色毛片三级朝国网站| 18+在线观看网站| 国产欧美日韩综合在线一区二区| 日本av手机在线免费观看| 国产成人欧美| 国产色婷婷99| 午夜精品国产一区二区电影| 在线观看一区二区三区激情| 免费大片黄手机在线观看| 欧美精品一区二区免费开放| 国精品久久久久久国模美| 午夜精品国产一区二区电影| 久久久国产精品麻豆| 亚洲综合色惰| 咕卡用的链子| 亚洲第一av免费看| 成人综合一区亚洲| 亚洲成人av在线免费| 国产精品国产三级国产av玫瑰| 国产精品熟女久久久久浪| 亚洲国产看品久久| 人成视频在线观看免费观看| 2018国产大陆天天弄谢| 春色校园在线视频观看| 亚洲国产看品久久| 99国产综合亚洲精品| 欧美bdsm另类| 青青草视频在线视频观看| 韩国av在线不卡| 久热这里只有精品99| 成人毛片a级毛片在线播放| 草草在线视频免费看| 黄片播放在线免费| 一边摸一边做爽爽视频免费| 日韩欧美一区视频在线观看| 美女大奶头黄色视频| 十分钟在线观看高清视频www| 成人无遮挡网站| 午夜福利视频在线观看免费| 日本免费在线观看一区| 18禁裸乳无遮挡动漫免费视频| 侵犯人妻中文字幕一二三四区| 色网站视频免费| 日本爱情动作片www.在线观看| 成年女人在线观看亚洲视频| 色婷婷av一区二区三区视频| 三上悠亚av全集在线观看| av网站免费在线观看视频| 午夜福利视频精品| 国产又色又爽无遮挡免| 五月玫瑰六月丁香| 亚洲少妇的诱惑av| 日韩伦理黄色片| 97人妻天天添夜夜摸| 亚洲成人av在线免费| 综合色丁香网| 日韩成人伦理影院| 2018国产大陆天天弄谢| tube8黄色片| 欧美人与性动交α欧美精品济南到 | 免费大片黄手机在线观看| 欧美另类一区| 国产xxxxx性猛交| 最新中文字幕久久久久| 国产一级毛片在线| 最新的欧美精品一区二区| 高清欧美精品videossex| 黄色配什么色好看| 两个人看的免费小视频| 免费看不卡的av| av又黄又爽大尺度在线免费看| 午夜精品国产一区二区电影| 少妇被粗大猛烈的视频| 999精品在线视频| 22中文网久久字幕| 亚洲av免费高清在线观看| 成人免费观看视频高清| 日韩成人av中文字幕在线观看| 欧美变态另类bdsm刘玥| 免费久久久久久久精品成人欧美视频 | 久久久a久久爽久久v久久| 一级片免费观看大全| 91精品国产国语对白视频| 极品少妇高潮喷水抽搐| 一区二区三区精品91| 午夜激情av网站| 免费女性裸体啪啪无遮挡网站| 国产精品久久久久久av不卡| 十八禁网站网址无遮挡| 欧美日韩视频高清一区二区三区二| 精品人妻在线不人妻| 免费不卡的大黄色大毛片视频在线观看| 亚洲美女搞黄在线观看| 91aial.com中文字幕在线观看| 欧美xxxx性猛交bbbb| av.在线天堂| av卡一久久| 欧美+日韩+精品| 免费av中文字幕在线| 午夜影院在线不卡| 日韩三级伦理在线观看| 国产精品麻豆人妻色哟哟久久| 国产一区二区在线观看av| 欧美成人精品欧美一级黄| 日产精品乱码卡一卡2卡三| 美女国产高潮福利片在线看| 日本黄大片高清| 热re99久久国产66热| 国产一区二区三区综合在线观看 | 人人妻人人澡人人爽人人夜夜| 国产成人av激情在线播放| 午夜免费男女啪啪视频观看| 精品久久国产蜜桃| 国产淫语在线视频| 有码 亚洲区| 一级爰片在线观看| 久久久国产一区二区| 日本欧美视频一区| 777米奇影视久久| 国产色爽女视频免费观看| 女人精品久久久久毛片| 国产亚洲av片在线观看秒播厂| 精品一区二区免费观看| 精品亚洲成a人片在线观看| 日本vs欧美在线观看视频| 国产深夜福利视频在线观看| 久久精品久久久久久噜噜老黄| 中文字幕制服av| 亚洲熟女精品中文字幕| 999精品在线视频| 亚洲色图 男人天堂 中文字幕 | 一级,二级,三级黄色视频| 下体分泌物呈黄色| 日韩人妻精品一区2区三区| 中文乱码字字幕精品一区二区三区| 天堂中文最新版在线下载| 2021少妇久久久久久久久久久| 欧美日韩一区二区视频在线观看视频在线| 99久国产av精品国产电影| 亚洲一码二码三码区别大吗| www日本在线高清视频| 国产免费福利视频在线观看| a 毛片基地| 精品人妻偷拍中文字幕| 男人爽女人下面视频在线观看| 国产免费一级a男人的天堂| 国产片特级美女逼逼视频| 边亲边吃奶的免费视频| 侵犯人妻中文字幕一二三四区| 亚洲精品乱码久久久久久按摩| 少妇 在线观看| 国产极品天堂在线| 国产精品久久久久久精品古装| 亚洲美女黄色视频免费看| 久久久精品区二区三区| 国产一区二区在线观看日韩| 国产爽快片一区二区三区| 一区二区av电影网| 十八禁高潮呻吟视频| 十分钟在线观看高清视频www| tube8黄色片| 欧美精品国产亚洲| 另类亚洲欧美激情| 人人妻人人添人人爽欧美一区卜| 精品酒店卫生间| 大片免费播放器 马上看| 大陆偷拍与自拍| 欧美人与性动交α欧美软件 | 超碰97精品在线观看| 国产 一区精品| 性色av一级| 五月伊人婷婷丁香| 男女午夜视频在线观看 | 国产探花极品一区二区| 国产免费视频播放在线视频| 狠狠婷婷综合久久久久久88av| 色婷婷久久久亚洲欧美| 丝袜人妻中文字幕| 三上悠亚av全集在线观看| 美女国产视频在线观看| h视频一区二区三区| 久久影院123| 99国产精品免费福利视频| 久久韩国三级中文字幕| 国产成人精品久久久久久| 午夜激情av网站| 国产精品偷伦视频观看了| 中文字幕制服av| 日日撸夜夜添| 亚洲精品久久成人aⅴ小说| 制服人妻中文乱码| 女人久久www免费人成看片| 国产在视频线精品| 日韩一本色道免费dvd| 啦啦啦啦在线视频资源| 极品人妻少妇av视频| 日本欧美视频一区| 两个人看的免费小视频| 亚洲人成77777在线视频| 国内精品宾馆在线| 亚洲精品日韩在线中文字幕| 午夜精品国产一区二区电影| 国产毛片在线视频| 日本爱情动作片www.在线观看| 人人妻人人澡人人爽人人夜夜| 99久久精品国产国产毛片| 久久免费观看电影| 美女视频免费永久观看网站| 亚洲国产av影院在线观看| 亚洲国产毛片av蜜桃av| 国产亚洲最大av| 国产在线视频一区二区| 亚洲综合精品二区| av网站免费在线观看视频| 欧美丝袜亚洲另类| 欧美人与性动交α欧美精品济南到 | 婷婷色麻豆天堂久久| 亚洲精品国产av成人精品| 亚洲国产精品一区二区三区在线| 一边摸一边做爽爽视频免费| 久久久久久久久久人人人人人人| 久久午夜福利片| 成人毛片a级毛片在线播放| 亚洲精品国产av成人精品| 成人毛片60女人毛片免费| 国产精品国产三级国产专区5o| 91午夜精品亚洲一区二区三区| 亚洲国产欧美日韩在线播放| 女人精品久久久久毛片| 在线观看人妻少妇| 国产视频首页在线观看| 九草在线视频观看| 欧美性感艳星| 免费在线观看完整版高清| 黑人欧美特级aaaaaa片| 欧美激情极品国产一区二区三区 | 亚洲欧美色中文字幕在线| 亚洲伊人久久精品综合| av线在线观看网站| 汤姆久久久久久久影院中文字幕| 欧美亚洲日本最大视频资源| 亚洲色图 男人天堂 中文字幕 | 看十八女毛片水多多多| 王馨瑶露胸无遮挡在线观看| 国产男女超爽视频在线观看| 丝袜脚勾引网站| 男的添女的下面高潮视频| 建设人人有责人人尽责人人享有的| 亚洲国产av新网站| 午夜免费观看性视频| 久久97久久精品| 寂寞人妻少妇视频99o| 免费日韩欧美在线观看| 国产一区二区三区av在线| 精品亚洲成国产av| av片东京热男人的天堂| 免费看不卡的av| 成人漫画全彩无遮挡| 日韩,欧美,国产一区二区三区| 综合色丁香网| 国产一区二区三区综合在线观看 | 成人免费观看视频高清| 国产片特级美女逼逼视频| 中文天堂在线官网| 久久久久久久国产电影| 亚洲美女视频黄频| 一区二区日韩欧美中文字幕 | 好男人视频免费观看在线| 国产免费一级a男人的天堂| 免费日韩欧美在线观看| 亚洲天堂av无毛| 亚洲欧美日韩另类电影网站| 大香蕉久久成人网| 久久久久精品性色| 日本色播在线视频| 晚上一个人看的免费电影| 爱豆传媒免费全集在线观看| 国产色婷婷99| 日日爽夜夜爽网站| 伊人亚洲综合成人网| 秋霞在线观看毛片| 少妇人妻久久综合中文| 一个人免费看片子| 色视频在线一区二区三区| 亚洲,欧美,日韩| 高清欧美精品videossex| 日本黄大片高清| 欧美成人精品欧美一级黄| 最新中文字幕久久久久| 久久精品aⅴ一区二区三区四区 | 九色成人免费人妻av| 欧美97在线视频| 亚洲,一卡二卡三卡| kizo精华| 咕卡用的链子| 大话2 男鬼变身卡| 纯流量卡能插随身wifi吗| 色哟哟·www| 久久av网站| 2021少妇久久久久久久久久久| 天天操日日干夜夜撸| 熟妇人妻不卡中文字幕| 国产视频首页在线观看| 国产av精品麻豆| 亚洲四区av| 亚洲天堂av无毛| 精品一品国产午夜福利视频| 久久国产亚洲av麻豆专区| 亚洲欧美一区二区三区黑人 | 男女无遮挡免费网站观看| 一区二区三区四区激情视频| 亚洲成人av在线免费| 国产亚洲一区二区精品| 成人国语在线视频| 欧美变态另类bdsm刘玥| av电影中文网址| 中文字幕另类日韩欧美亚洲嫩草| 久热久热在线精品观看| 国产男女内射视频| 人妻 亚洲 视频| 国产免费一区二区三区四区乱码| 亚洲第一区二区三区不卡| 女人被躁到高潮嗷嗷叫费观| 九色成人免费人妻av| 高清视频免费观看一区二区| 欧美 亚洲 国产 日韩一| 亚洲天堂av无毛| 精品亚洲成a人片在线观看| 午夜免费鲁丝| 波野结衣二区三区在线| 性色av一级| 亚洲国产欧美在线一区| 一区二区日韩欧美中文字幕 | 少妇精品久久久久久久| 日韩精品有码人妻一区| 久久国产精品男人的天堂亚洲 | 在线观看人妻少妇| 欧美另类一区| 看十八女毛片水多多多| 99热这里只有是精品在线观看| 国产男人的电影天堂91| 日韩欧美一区视频在线观看| 国产av国产精品国产| 91午夜精品亚洲一区二区三区| 一边摸一边做爽爽视频免费| 女人精品久久久久毛片| 成年人免费黄色播放视频| 成人漫画全彩无遮挡| 男女下面插进去视频免费观看 | 日韩中字成人| 80岁老熟妇乱子伦牲交| 我要看黄色一级片免费的| 亚洲av中文av极速乱| 国产日韩欧美在线精品| 亚洲成人手机| 亚洲国产精品一区二区三区在线| 高清毛片免费看| 天堂8中文在线网| 人体艺术视频欧美日本| 欧美 日韩 精品 国产| 国产视频首页在线观看| 久久久久久久久久久免费av| 久久99热6这里只有精品|