Performance of induction module cavity ring-down spectroscopy (IM-CRDS) for measuring δ18 O and δ2 H values of soil, stem, and leaf waters.

RATIONALE: Induction module cavity ring-down spectroscopy (IM-CRDS) has been proposed as a rapid and cost-effective alternative to cryogenic vacuum distillation (CVD) and isotope ratio mass spectrometry (IRMS) for the measurement of δ18 O and δ2 H values in matrix-bound waters. In the current study, we characterized the performance of IM-CRDS relative to CVD and IRMS and investigated the mechanisms responsible for differences between the methods. METHODS: We collected a set of 75 soil, stem, and leaf water samples, and measured the δ18 O and δ2 H values of each sample with four techniques: CVD and IRMS, CVD and CRDS, CVD and IM-CRDS, and IM-CRDS alone. We then calculated the isotopic errors for each of the three CRDS methods relative to CVD and IRMS, and analyzed the relationships among these errors and suites of diagnostic spectral parameters that are indicative of organic contamination. RESULTS: The IM-CRDS technique accurately assessed the δ18 O and δ2 H values of pure waters, but exhibited progressively increasing errors for soil waters, stem waters, and leaf waters. For soils, the errors were attributable to subsampling of isotopically heterogeneous source material, whereas for stems and leaves, they were attributable to spectral interference. Unexpectedly, the magnitude of spectral interference was higher for the solid samples analyzed directly via IM-CRDS than for those originally extracted via CVD and then analyzed by IM-CRDS. CONCLUSIONS: There are many types of matrix-bound water samples for which IM-CRDS measurements include significant errors from spectral interference. As a result, spectral analysis and validation should be incorporated into IM-CRDS post-processing procedures. In the future, IM-CRDS performance could be improved through: (i) identification of the compounds that cause spectral interference, and either (ii) modification of the combustion step to completely oxidize these compounds to CO2 , and/or (iii) incorporation of corrections for these compounds into the spectral fitting models used by the CRDS analyzers. Copyright © 2016 John Wiley & Sons, Ltd.

Herbivory-induced mortality increases with radial growth in an invasive riparian phreatophyte.

BACKGROUND AND AIMS: Under equal conditions, plants that allocate a larger proportion of resources to growth must do so at the expense of investing fewer resources to storage. The critical balance between growth and storage leads to the hypothesis that in high-resource environments, plants that express high growth rates are more susceptible to episodic disturbance than plants that express lower growth rates. METHODS: This hypothesis was tested by measuring the radial growth, basal area increment (BAI) and carbon isotope ratios (δ(13)C) in tree-ring α-cellulose of 62 mature tamarisk trees (Tamarix spp.) occurring at three sites in the western USA (n = 31 live and 31 killed trees across all sites, respectively). All of the trees had been subjected to periods of complete foliage loss by episodic herbivory over three or more consecutive growing seasons by the tamarisk leaf beetle (Diorhabda carinulata), resulting in approx. 50 % mortality at each site. KEY RESULTS: Mean annual BAI (measured from annual ring widths) in the 10 years prior to the onset of herbivory was on average 45 % higher in killed trees compared with live trees (P < 0·0001). Killed trees that had higher growth rates also expressed higher (less negative) δ(13)C ratios compared with live trees. In fact, at one site near Moab, UT, the mean annual BAI was 100 % higher in killed trees despite having about a 0·5 ‰ higher δ(13)C relative to live trees (P = 0·0008). Patterns of δ(13)C suggest that the intrinsic water-use efficiency was higher in killed than surviving trees, possibly as a consequence of lower whole-canopy stomatal conductance relative to live trees. CONCLUSIONS: The results show that a likely trade-off occurs between radial growth and survival from foliage herbivory in Tamarix spp. that currently dominates riparian areas throughout the western USA and northern Mexico. Thus, herbivory by D. carinulata may reduce the overall net primary productivity of surviving Tamarix trees and may result in a reduction in genetic variability in this dominant invasive tree species if these allocation patterns are adaptive.

Trends in Stomatal Density and 13C/12C Ratios of Pinus flexilis Needles During Last Glacial-Interglacial Cycle.

Measurements of stomatal density and delta(13)C of limber pine (Pinus flexilis) needles (leaves) preserved in pack rat middens from the Great Basin reveal shifts in plant physiology and leaf morphology during the last 30,000 years. Sites were selected so as to offset glacial to Holocene climatic differences and thus to isolate the effects of changing atmospheric CO(2) levels. Stomatal density decreased approximately 17 percent and delta(13)C decreased approximately 1.5 per mil during deglaciation from 15,000 to 12,000 years ago, concomitant with a 30 percent increase in atmospheric CO(2). Water-use efficiency increased approximately 15 percent during deglaciation, if temperature and humidity were held constant and the proxy values for CO(2) and delta(13)C of past atmospheres are accurate. The delta(13)C variations may help constrain hypotheses about the redistribution of carbon between the atmosphere and biosphere during the last glacial-interglacial cycle.

Acclimation response of spring wheat in a free-air CO(2) enrichment (FACE) atmosphere with variable soil nitrogen regimes. 1. Leaf position and phenology determine acclimation response.

We have examined the photosynthetic acclimation of wheat leaves grown at an elevated CO(2) concentration, and ample and limiting N supplies, within a field experiment using free-air CO(2) enrichment (FACE). To understand how leaf age and developmental stage affected any acclimation response, measurements were made on a vertical profile of leaves every week from tillering until maturity. The response of assimilation (A) to internal CO(2) concentration (C(i)) was used to estimate the in vivo carboxylation capacity (Vc(max)) and maximum rate of ribulose-1,5-bisphosphate limited photosynthesis (A (sat)). The total activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and leaf content of Rubisco and the Light Harvesting Chlorophyll a/b protein associated with Photosystem II (LHC II), were determined. Elevated CO(2) did not alter Vc(max) in the flag leaf at either low or high N. In the older shaded leaves lower in the canopy, acclimatory decline in Vc(max) and A (sat) was observed, and was found to correlate with reduced Rubisco activity and content. The dependency of acclimation on N supply was different at each developmental stage. With adequate N supply, acclimation to elevated CO(2) was also accompanied by an increased LHC II/Rubisco ratio. At low N supply, contents of Rubisco and LHC II were reduced in all leaves, although an increased LHC II/Rubisco ratio under elevated CO(2) was still observed. These results underscore the importance of leaf position, leaf age and crop developmental stage in understanding the acclimation of photosynthesis to elevated CO(2) and nutrient stress.

Acclimation response of spring wheat in a free-air CO(2) enrichment (FACE) atmosphere with variable soil nitrogen regimes. 3. Canopy architecture and gas exchange.

The response of whole-canopy net CO(2) exchange rate (CER) and canopy architecture to CO(2) enrichment and N stress during 1996 and 1997 for open-field-grown wheat ecosystem (Triticum aestivum L. cv. Yecora Rojo) are described. Every Control (C) and FACE (F) CO(2) treatment (defined as ambient and ambient +200 mumol mol(-1), respectively) contained a Low- and High-N treatment. Low-N treatments constituted initial soil content amended with supplemental nitrogen applied at a rate of 70 kg N ha(-1) (1996) and 15 kg N ha(-1) (1997), whereas High-N treatments were supplemented with 350 kg N ha(-1) (1996 and 1997). Elevated CO(2) enhanced season-long carbon accumulation by 8% and 16% under Low-N and High-N, respectively. N-stress reduced season-long carbon accumulation 14% under ambient CO(2), but by as much as 22% under CO(2) enrichment. Averaging both years, green plant area index (GPAI) peaked approximately 76 days after planting at 7.13 for FH, 6.00 for CH, 3.89 for FL, and 3.89 for CL treatments. Leaf tip angle distribution (LTA) indicated that Low-N canopies were more erectophile than those of High-N canopies: 48 degrees for FH, 52 degrees for CH, and 58 degrees for both FL and CL treatments. Temporal trends in canopy greenness indicated a decrease in leaf chlorophyll content from the flag to flag-2 leaves of 25% for FH, 28% for CH, 17% for CL, and 33% for FL during 1997. These results indicate that significant modifications of canopy architecture occurs in response to both CO(2) and N-stress. Optimization of canopy architecture may serve as a mechanism to diminish CO(2) and N-stress effects on CER.

Acclimation response of spring wheat in a free-air CO(2) enrichment (FACE) atmosphere with variable soil nitrogen regimes. 2. Net assimilation and stomatal conductance of leaves.

Atmospheric CO(2) concentration continues to rise. It is important, therefore, to determine what acclimatory c hanges will occur within the photosynthetic apparatus of wheat (Triticum aestivum L. cv. Yecora Rojo) grown in a future high-CO(2) world at ample and limited soil N contents. Wheat was grown in an open field exposed to the CO(2) concentration of ambient air [370 mumol (CO(2)) mol(-1); Control] and air enriched to approximately 200 mumol (CO(2)) mol(-1) above ambient using a Free-Air CO(2) Enrichment (FACE) apparatus (main plot). A High (35 g m(-2)) or Low (7 and 1.5 g m(-2) for 1996 and 1997, respectfully) level of N was applied to each half of the main CO(2) treatment plots (split-plot). Under High-N, FACE reduced stomatal conductance (g (s)) by 30% at mid-morning (2 h prior to solar noon), 36% at midday (solar noon) and 27% at mid-afternoon (2.5 h after solar noon), whereas under Low-N, g (s) was reduced by as much as 31% at mid-morning, 44% at midday and 28% at mid-afternoon compared with Control. But, no significant CO(2) x N interaction effects occurred. Across seasons and growth stages, daily accumulation of carbon (A') was 27% greater in FACE than Control. High-N increased A' by 18% compared with Low-N. In contrast to results for g (s), however, significant CO(2) x N interaction effects occurred because FACE increased A' by 30% at High-N, but by only 23% at Low-N. FACE enhanced the seasonal accumulation of carbon (A'') by 29% during 1996 (moderate N-stress), but by only 21% during 1997 (severe N-stress). These results support the premise that in a future high-CO(2) world an acclimatory (down-regulation) response in the photosynthetic apparatus of field-grown wheat is anticipated. They also demonstrate, however, that the stimulatory effect of a rise in atmospheric CO(2) on carbon gain in wheat can be maintained if nutrients such as nitrogen are in ample supply.

The effect of long-term atmospheric CO2 enrichment on the intrinsic water-use efficiency of sour orange trees.

Every two months of 1992, as well as on three occasions in 1994-1995, we obtained leaf samples together with samples of surrounding air from eight well-watered and fertilized sour orange (Citrus aurantium L.) trees that were growing out-of-doors at Phoenix, Arizona, USA. These trees had been planted in the ground as small seedlings in July of 1987 and enclosed in pairs by four clear-plastic-wall open-top chambers of which two have been continuously maintained since November of that year at a CO2 concentration of 400 micromol mol(-1) and two have been maintained at 700 micromol mol(-1). In September 2000, we also extracted north-south and east-west oriented wood cores that passed through the center of each tree's trunk at a height of 45 cm above the ground. Stable-carbon isotope ratios (13C/12C) derived from these leaf, wood and air samples were used to evaluate each tree's intrinsic water-use efficiency (iWUE). The grand-average result was an 80% increase in this important plant parameter in response to the 300 micromol mol(-1) increase in atmospheric CO2 concentration employed in the study. This increase in sour orange tree iWUE is identical to the long-term CO2-induced increase in the trees' production of wood and fruit biomass, which suggests there could be little to no change in total water-use per unit land area for this species as the air's CO2 content continues to rise. It is also identical to the increase in the mean iWUE reported for 23 groups of naturally occurring trees scattered across western North America that was caused by the historical rise in the air's CO2 content that occurred between 1800 and 1985.

Stable-carbon isotopic composition of maple sap and foliage.

The (13)C/(12)C ratios of Acer grandidentatum sap sugar collected during the dormant period are compared to those of buds, leaves, and wood developed over the following growing season. As the primary carbon source for cellulose manufacture at initiation of annual growth in deciduous trees, sap sucrose would be expected to have an isotopic composition similar to first-formed cellulose. Although constancy in concentration and (13)C/(12)C ratios of the maple sap sugar suggests any gains or losses (e.g. to maintenance metabolism) do not appreciably alter composition, the (13)C/(12)C ratios of cellulose of the enlarging buds in the spring are quite distinct from those of the sap sugar, seemingly precluding a simple direct biochemical pathway of sap sucrose-->glucose-->cellulose in favor of a more complex pathway with greater likelihood of isotopic fractionation. The (13)C/(12)C ratios of the leaves and in the growth ring were initially similar to the sap sugar but decreased steadily over the growing season.

Development of C4 photosynthesis in sorghum leaves grown under free-air CO2 enrichment (FACE).

The developmental pattern of C4 expression has been well characterized in maize and other C4 plants. However, few reports have explored the possibility that the development of this pathway may be sensitive to changes in atmospheric CO2 concentrations. Therefore, both the structural and biochemical development of leaf tissue in the fifth leaf of Sorghum bicolor plants grown at elevated CO2 have been characterized. Ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC) activities accumulate rapidly as the leaf tissue differentiates and emerges from the surrounding whorl. Rubisco was not expressed in a cell-specific manner in the youngest tissue at the base of the leaf, but did accumulate before PEPC was detected. This suggests that the youngest leaf tissue utilizes a C3-like pathway for carbon fixation. However, this tissue was in a region of the leaf receiving very low light and so significant rates of photosynthesis were not likely. Older leaf tissue that had emerged from the surrounding whorl into full sunlight showed the normal C4 syndrome. Elevated CO2 had no effect on the cell-specific localization of Rubisco or PEPC at any stage of leaf development, and the relative ratios of Rubisco to PEPC remained constant during leaf development. However, in the oldest tissue at the tip of the leaf, the total activities of Rubisco and PEPC were decreased under elevated CO2 implying that C4 photosynthetic tissue may acclimate to growth under elevated CO2.