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Elevated atmospheric CO2 and drought effects on leaf gas exchange properties of barley
Authors:Gerard W Wall  Richard L GarciaFrank Wechsung  Bruce A Kimball
Institution:a U.S. Arid-Land Agricultural Research Center, USDA-ARS, 21881 North Cardon Lane, Maricopa, AZ 85238, USA
b LI-COR BioScience Inc., P.O. Box 4425, Lincoln, NE 68504, USA
c Potsdam-Institute for Climate Impact Research, P.O. Box 601203, D-14412 Potsdam, Germany
Abstract:Atmospheric CO2 concentration (Ca) is rising, predicted to cause global warming, and alter precipitation patterns. During 1994, spring barley (Hordeum vulgare L. cv. Alexis) was grown in a strip-split-plot experimental design to determine the effects that the main plot Ca treatments A: Ambient at 370 μmol (CO2) mol−1; E: Enriched with free-air CO2 enrichment (FACE) at ∼550 μmol (CO2) mol−1] had on several gas exchange properties of fully expanded sunlit primary leaves. The interacting strip-split-plot irrigation treatments were Dry or Wet 50% (D) or 100% (W) replacement of potential evapotranspiration] at ample nitrogen (261 kg N ha−1) and phosphorous (29 kg P ha−1) fertility. Elevated Ca facilitated drought avoidance by reducing stomatal conductance (gs) by 34% that conserved water and enabled stomata to remain open for a longer period into a drought. This resulted in a 28% reduction in drought-induced midafternoon depression in net assimilation rate (A). Elevated Ca increased A by 37% under Dry and 23% under Wet. Any reduction in A under Wet conditions occurred because of nonstomatal limitations, whereas under Dry it occurred because of stomatal limitations. Elevated Ca increased the diurnal integral of A (A′) that resulted in an increase in the seasonal-long integral of A′ (A″) for barley leaves by 12% (P = 0.14) under both Dry and Wet - 650, 730, 905 and 1020 ± 65 g (C) m−2 y−1 for AD, ED, AW and EW treatments, respectively. Elevated Ca increased season-long average dry weight (DWS; crown, shoots) by 14% (P = 0.02), whereas deficit irrigation reduced DWS by 7% (P = 0.06), although these values may have been affected by a short but severe pea aphid Acyrthosiphon pisum (Harris)] infestation. Hence, an elevated-Ca-based improvement in gas exchange properties enhanced growth of a barley crop.
Keywords:A  Ambient Ca treatment  A  instantaneous leaf net assimilation rate [μmol (CO2) m&minus    s&minus  1]  Amax  daily maximum leaf net assimilation rate [μmol (CO2) m&minus    s&minus  1]  AN  season-long relative mean leaf net assimilation rate normalized between 0 and 1 scale (i  e    AN     A/Amax  dimensionless)  A&prime    daily integral of net leaf carbon accumulation [g (C) m&minus    d&minus  1]  A&Prime    seasonal integral of net leaf carbon accumulation [g (C) m&minus    y&minus  1]  AD  Ambient-Dry treatment  ANOVA  analysis of variance effects (i  e    C  D  I  S  T)  AW  Ambient-Wet treatment  df1  first degrees of freedom for F-statistic  df2  second degrees of freedom for F-statistic  C  carbon dioxide effect in ANOVA  Ca  atmospheric CO2 concentration [μmol (CO2  mol&minus  1]  Ci  intercellular CO2 concentration [μmol (CO2) mol&minus  1]  Ci/Ca  ratio of Ci to Ca (dimensionless)  D  Dry irrigation treatment  D  soil dehydration cycle effect in ANOVA  DAE  day after 50% emergence  DWG  dry weight of grain for barley (g   plant&minus  1)  DWS  dry weight of shoot and crown tissue for barley (g   plant&minus  1)  DWS  B:W  percentage change in dry weight between barley and wheat (%)  DWS  W  dry weight of shoot and crown tissue for wheat (g   plant&minus  1)  E  Enriched Ca treatment  E  leaf transpiration rate [mmol (H2O) m&minus    s&minus  1]  ED  Enriched-Dry treatment  EW  Enriched-Wet treatment  ea  atmospheric water vapor pressure at Ta (Pa)  e*  atmospheric saturation water vapor pressure at Ta (Pa)  ea  VPD  atmospheric water vapor pressure deficit (i  e    ea  VPD     e*   &minus     ea) at Ta (Pa)  F  F-statistic  FACE  free-air-CO2-enrichment  gs  stomatal conductance to water vapor [mol (H2O) m&minus    s&minus  1]  I  irrigation effect in ANOVA  IWUE (A/gs)  intrinsic water use efficiency [μmol (CO2) mol (H2O)&minus  1]  PPFD  photosynthetic photon flux density [μmol (photons) m&minus    s&minus  1]  Pα  probability of a greater F-statistic by chance  S  species effect in ANOVA comparing average season-long dry weight response of barley to wheat  SR  daily total global solar radiation (MJ   m&minus    d&minus  1)  SRi  instantaneous total global solar radiation (MJ   m&minus    h&minus  1)  T  time of day [midmorning (MM: 2    h prior to solar noon)  midday (MD: solar noon)  and midafternoon (MA: 2    h after solar noon)] effect in ANOVA  TDR  time domain reflectometry  Ta  ambient air temperature inside cuvette (°  C)  Tl  leaf temperature inside cuvette (°  C)  Ta  max  maximum Ta  C)  Ta  min  minimum Ta  C)  TS  soil temperature (°  C)  TS  max  maximum TS  C)  TS  min  minimum TS  C)  ΔT  leaf minus air temperature inside cuvette (°  C)  ua  avg  average annual daily wind speed (m   s&minus  1) at 2-m height  W  Wet irrigation treatment  WUE (A/E)  water use efficiency [μmol (CO2) mmol (H2O)&minus  1]  ΨM  soil matric potential (MPa)  θS  volumetric soil-water content (m3   m&minus  3)
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