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Consequences of elevated carbon dioxide and ozone for foliar chemical composition and dynamics in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera)

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Abstract

Atmospheric chemical composition affects foliar chemical composition, which in turn influences the dynamics of both herbivory and decomposition in ecosystems. We assessed the independent and interactive effects of CO2 and O3 fumigation on foliar chemistry of quaking aspen (Populus tremuloides) and paper birch (Betula papyrifera) at a Free-Air CO2 Enrichment (FACE) facility in northern Wisconsin. Leaf samples were collected at five time periods during a single growing season, and analyzed for nitrogen, starch and condensed tannin concentrations, nitrogen resorption efficiencies (NREs), and C:N ratios. Enriched CO2 reduced foliar nitrogen concentrations in aspen and birch; O3 only marginally reduced nitrogen concentrations. NREs were unaffected by pollution treatment in aspen, declined with O3 exposure in birch, and this decline was ameliorated by enriched CO2. C:N ratios of abscised leaves increased in response to enriched CO2 in both tree species. O3 did not significantly alter C:N ratios in aspen, although values tended to be higher in +CO2+O3 leaves. For birch, O3 decreased C:N ratios under ambient CO2 and increased C:N ratios under elevated CO2. Thus, under the combined pollutants, the C:N ratios of both aspen and birch leaves were elevated above the averaged responses to the individual and independent trace gas treatments. Starch concentrations were largely unresponsive to CO2 and O3 treatments in aspen, but increased in response to elevated CO2 in birch. Levels of condensed tannins were negligibly affected by CO2 and O3 treatments in aspen, but increased in response to enriched CO2 in birch. Results from this work suggest that changes in foliar chemical composition elicited by enriched CO2 are likely to impact herbivory and decomposition, whereas the effects of O3 are likely to be minor, except in cases where they influence plant response to CO2.

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... Tropospheric ozone has the capacity to impact on nutrient cycling by both direct and indirect mechanisms, namely by (1) altering the chemical composition of plant tissue, (2) altering the quantity (and quality) of litter fall, (3) impacting on below-ground plant biomass and root exudates, (4) indirectly altering microbial community composition(s) and functioning, and (5) indirectly influencing soil processes and the chemical properties of soils. All of these have the capacity either, independently or in concert, to reduce the long-term sustainability of ecosystems (Lindroth et al., 2001). ...
... Reduced N uptake under 2 x ambient ozone has been previously found in beech by Haberer and colleagues (2007). Lui et al. (2005) and Lindroth et al. (2001) also report significantly reduced foliar N by up to 6.6% in response to elevated ozone. ...
... The effects of ozone on secondary metabolites, lignifications and/or carbon: nitrogen (C: N) ratio of above-and below-ground plant parts can also alter the biodegradability of litter and thus the rate of decomposition which may, in turn, impact on detritivore and herbivore trophic pathways (Lindroth et al., 2001). Boerner and Rebbeck, (1995) have shown that ozone altered the structure, nutrient status and deposition of secondary metabolites in leaves of forest trees, whereby ozone lowered soluble carbohydrate concentrations and increased lignin content, the consequences of which were reduced decay rates of litter fall. ...
Technical Report
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In this report we provide a review of the state of current knowledge on the effects of ozone pollution on ecosystem services including consideration of effects on biodiversity. Although considered separately, all of the ecosystem services and underlying processes are interlinked, with for example, ozone impacts on root growth contributing to supporting services (primary productivity), provisioning (crop and timber production), regulating (C sequestration and impacts on climate) and cultural services (reduced growth of sensitive species influencing aesthetic qualities of vegetation), and reducing economic value of products such as crop yield. Until recently, much of the research on ozone impacts has focussed on quantifying effects on ecological processes rather than considering the implications for ecosystem services. This report, for the first time, places current process-based knowledge within the context of ecosystem services and thus reports on the potential for impacts of ozone on ecosystem services and biodiversity. The report has been prepared by the Coordination Centre of the ICP Vegetation, an International Cooperative Programme reporting on air pollution impacts on vegetation to the Working Group on Effects of the Convention on Long-Range Transboundary Air Pollution.
... Their results and ours are consistent with a modeling study (Chen et al. 1994) which predicted that ozone might have little effects on plant biomass because of compensatory regrowth of more efficient younger tissues when ozone accelerated senescence. Lindroth et al. (2001) found that foliar C:N of birch and aspen had non-additive responses to elevated ozone and CO 2 with large increases in C:N in combination. In addition, it is possible that soil microorganisms could have non-additive responses that shape plant responses which is consistent with mass being highest on average in CO 2 + O 3 for every species but we have no data to evaluate this possibility. ...
... The effects of ozone and CO 2 on condensed tannins were minor for most of the species in this study, including T. sebifera. Lindroth et al. (2001) found that birch and aspen also 1: Interactive effects of ozone and CO 2 on leaf, stem, root, and total mass and leaf number and root:shoot. The means for O 3 , CO 2 and CO 2 + O 3 are shown as ratios relative to control means. ...
... This is most similar to the results we observed for L. formosana (China) that had greater tannin concentrations in CO 2 + O 3 than in other treatments. However, Lindroth et al. (2001) tested plants that had been exposed for >1 year which may limit comparison to our study in which plants were exposed for ~11 weeks. Moreover, the results for growth and secondary metabolites may be related as secondary metabolites are considered an effective mechanism to defend against environmental stresses including abiotic and biotic factors (Bartwal et al. 2013). ...
Article
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Aims: Elevated ozone and CO2 can differentially affect the performance of plant species. Variation among native, exotic, and invader species in their growth and defense responses to CO2 and ozone may shape CO2 and ozone effects on invasions, perhaps in part also due to variation between native and invasive populations of invaders. Methods: We manipulated ozone (control or 100 ppb) and CO2 (ambient or 800 ppm) in a factorial greenhouse experiment in replicated chambers. We investigated growth and defense (tannins) of seedlings of Triadica sebifera from invasive (US) and native (China) populations and pairs of US and China tree species within three genera (Celtis, Liquidambar, Platanus). Important findings: Overall, ozone reduced growth in ambient CO2 but elevated CO2 limited this effect. Triadica sebifera plants from invasive populations had higher growth than those from native populations in control conditions or the combination of elevated CO2 and ozone in which invasive populations had greater increases in growth. Their performances were similar in elevated CO2 because native populations were more responsive and their performances were similar with elevated ozone because invasive populations were more susceptible. Compared to other species, T. sebifera had high growth rates but low levels of tannin production that were insensitive to variation in CO2 or ozone. Both China and US Platanus plants reduced tannins with increased CO2 and/or ozone and US Liquidambar plants increased tannins with the combination of elevated CO2 and ozone. The growth results suggest that intraspecific variation in T. sebifera will reduce the effects of CO2 or ozone alone on invasions but increase their combined effects. The tannin results suggest that defense responses to CO2 and ozone will be variable across native and exotic species. The effects of CO2 and ozone on growth and defense of native and exotic species indicate that the benefit or harm to species from these global change drivers is an idiosyncratic combination of species origin and genus.
... As plant growth becomes limited by factors other than photosynthate supply (e.g., mineral nutrients), an increasing proportion of photosynthetic production becomes available for the synthesis of secondary compounds (Herms The effect is considered significant if the 95% CI does not overlap 1. Sources of data: , Booker and Miller 1998, Gebauer et al. 1998, Heyworth et al. 1998, Kainulainen et al. 1998, Booker and Maier 2001, King et al. 2001a, 2001c, Lindroth et al. 2001, Sallas et al. 2001, Peñuelas et al. 2002, Holton et al. 2003, Chapman 2004, Parsons et al. 2004and Booker et al. 2005 and Mattson 1992). We also hypothesized that the decline in photosynthesis in elevated [O 3 ] (Anderson 2003) reduces the availability of photosynthate for C-based defensive compounds. ...
... Booker (2000) reported that elevated [O 3 ] increased foliar [N] in cotton, whereas Scherzer et al. (1998) found that elevated [O 3 ] had no significant effect on foliar [N] of yellow-poplar or white pine. In the earlier study at the Aspen FACE, Parsons et al. (2004) reported that elevated [O 3 ] had no effect on paper birch litter [N], but Lindroth et al. (2001) found that elevated [O 3 ] significantly reduced aspen foliar [N]. In our study, elevated [O 3 ] appeared to reduce mean litter [N] by 6.6% (P = 0.073), and significantly increased the litter C:N ratio (P = 0.043). ...
... The effect is considered significant if the 95% CI does not overlap 1. Sources of data: Kull et al. 1996, Pfirrmann et al. 1996, Scherzer et al. 1998, Booker 2000, Utriainen et al. 2000, Lindroth et al. 2001, Parsons et al. 2004and Booker et al. 2005 We predicted that the phytotoxic effect of elevated [O 3 ] would lower concentrations of secondary compounds as a result of a reduction in carbohydrate supply. However, we found that elevated [O 3 ] significantly increased leaf litter concentrations of soluble sugars, soluble phenolics and condensed tannins, indicating that elevated [O 3 ] may trigger a biochemical defense or damage response that elevates synthesis of secondary compounds despite lower carbohydrate availability. ...
Article
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Human activities are increasing the concentrations of atmospheric carbon dioxide ([CO2]) and tropospheric ozone ([O-3]), potentially leading to changes in the quantity and chemical quality of leaf litter inputs to forest soils. Because the quality and quantity of labile and recalcitrant carbon (C) compounds influence forest productivity through changes in soil organic matter content, characterizing changes in leaf litter in response to environmental change is critical to understanding the effects of global change on forests. We assessed the independent and combined effects of elevated [CO2] and elevated [O-3] on foliar litter production and chemistry in aspen (Populus tremuloides Michx.) and birch-(Betula papyrifera Marsh.) aspen communities at the Aspen free-air CO2 enrichment (FACE) experiment in Rhinelander, WI. Litter was analyzed for concentrations of C, nitrogen (N), soluble sugars, lipids, lignin, cellulose, hemicellulose and C-based defensive compounds (soluble phenolics and condensed tannins). Concentrations of these chemical compounds in naturally senesced litter were similar in aspen and birch-aspen communities among treatments, except for N, the C:N ratio and lipids. Elevated [CO2] significantly increased C:N (+8.7%), lowered mean litter N concentration (-10.7%) but had no effect on the concentrations of soluble sugars, soluble phenolics and condensed tannins. Elevated [CO2] significantly increased litter biomass production (+33.3%), resulting in significant increases in fluxes of N, soluble sugars, soluble phenolics and condensed tannins to the soil. Elevated [O-3] significantly increased litter concentrations of soluble sugars (+78.1%), soluble phenolics (+53.1%) and condensed tannins (+77.2%). There were no significant effects of elevated [CO2] or elevated [O-3] on the concentrations of individual C structural carbohydrates (cellulose, hemicellulose and lignin). Elevated [CO2] significantly increased cellulose (+37.4%) input to soil, whereas elevated [O-3] significantly reduced hemicellulose and lignin inputs to soil (-22.3 and -31.5%, respectively). The small changes in litter chemistry in response to elevated [CO2] and tropospheric [O-3] that we observed, combined with changes in litter biomass production, could significantly alter the inputs of N, soluble sugars, condensed tannins, soluble phenolics, cellulose and lignin to forest soils in the future.
... The limited effect of CO 2 on foliar nitrogen contrasts with the reviews cited above and also with reports from earlier studies at Aspen FACE (e.g., Kopper and Lindroth 2003;Lindroth et al. 2001Lindroth et al. , 2002. Our findings, however, are consistent with multiple studies, including more recent ones at this research site, that report minimal effects of elevated CO 2 Fig. 1 Principal components (MDS) ordination plots illustrating differences between composite chemical profiles of aspen and birch leaves exposed the independent and interactive effects of ambient and elevated CO 2 and O 3 . ...
... Foliar quality data used to create both plots were averaged across genotypes and time for aspen and across time for birch on foliar nitrogen (e.g., Vigue and Lindroth 2010). Our results also agree with reports from Aspen FACE that found nominal effects of CO 2 on carbohydrate concentrations (e.g., Lindroth et al. 2001Lindroth et al. , 2002. ...
Article
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Anthropogenic activities are altering levels of atmospheric carbon dioxide (CO2) and tropospheric ozone (O3). These changes can alter phytochemistry, and in turn, influence ecosystem processes. We assessed the individual and combined effects of elevated CO2 and O3 on the phytochemical composition of two tree species common to early successional, northern temperate forests. Trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) were grown at the Aspen FACE (Free-Air Carbon dioxide and ozone Enrichment) facility under four combinations of ambient and elevated CO2 and O3. We measured, over three years (2006–08), the effects of CO2 and O3 on a suite of foliar traits known to influence forest functioning. Elevated CO2 had minimal effect on foliar nitrogen and carbohydrate levels in either tree species, and increased synthesis of condensed tannins and fiber in aspen, but not birch. Elevated O3 decreased nitrogen levels in both tree species and increased production of sugar, condensed tannins, fiber, and lignin in aspen, but not birch. The magnitude of responses to elevated CO2 and O3 varied seasonally for both tree species. When co-occurring, CO2 offset most of the changes in foliar chemistry expressed under elevated O3 alone. Our results suggest that levels of CO2 and O3 predicted for the mid-twenty-first century will alter the foliar chemistry of northern temperate forests with likely consequences for forest community and ecosystem dynamics.
... The decrease in leaf N concentration under O 3 exposure, as observed here, has also been found in other studies (Lindroth et al. 2001, Yamaji et al. 2003, Kopper and Lindroth 2003, Ribas et al. 2005, and is linked to O 3 -induced activation of senescence-related processes (Bielenberg et al. 2002). The decline in leaf N level resulted in a higher leaf C:N ratio, since leaf C concentrations remained unaffected under elevated O 3 . ...
... The decline in leaf N level resulted in a higher leaf C:N ratio, since leaf C concentrations remained unaffected under elevated O 3 . Similar to the findings of Lindroth et al. (2001) on trembling aspen, we did not find any influence of O 3 on the N resorption of the trees. Thus the N levels in leaf litter reflected that of green leaves, which contributed to the trend towards a higher C:N ratio in the European aspen litter that was produced under elevated O 3 . ...
Article
We studied growth and foliar responses of two clones of both European aspen (Populustremula) and hybrid aspen (P. tremula × P. tremuloides) to elevated O3 (45 ppb, 14-h mean) over one growing season using a free-air fumigation system in central Finland. All clones exhibited O3-specific foliar injury and accelerated leaf senescence under elevated O3. Yet, exposure to 1.5 × ambient O3 had only minor effects on the growth and biomass production of clones grown under optimal nutrient and water supply, and no O3 effects on leaf morphology were observed. Slower-growing European aspen was more sensitive to elevated O3 than hybrid aspen. Exposure to O3 decreased the root/stem ratio (-11%) and leaf N concentration (-9%) of European aspen. Inter- or intraspecific differences in the O3 sensitivity of the trees could not be explained by stomatal conductance, but some xeromorphic leaf traits were related to increased susceptibility to O3. Intraspecific differences in the O3 sensitivity have implications e.g. for nurseries producing commercial tree material.
... The decrease in leaf N concentration under O 3 exposure, as observed here, has also been found in other studies (Lindroth et al. 2001, Yamaji et al. 2003, Kopper and Lindroth 2003, Ribas et al. 2005, and is linked to O 3 -induced activation of senescence-related processes (Bielenberg et al. 2002). The decline in leaf N level resulted in a higher leaf C:N ratio, since leaf C concentrations remained unaffected under elevated O 3 . ...
... The decline in leaf N level resulted in a higher leaf C:N ratio, since leaf C concentrations remained unaffected under elevated O 3 . Similar to the findings of Lindroth et al. (2001) on trembling aspen, we did not find any influence of O 3 on the N resorption of the trees. Thus the N levels in leaf litter reflected that of green leaves, which contributed to the trend towards a higher C:N ratio in the European aspen litter that was produced under elevated O 3 . ...
... Evaluation of O 3 -induced impacts on nutrient translocation in plants from older to newer leaves has revealed an early shift in the micro-and macronutrients content in the leaf litter, at the outset of the stress exposure, which influences the ecological stoichiometry (Shi et al. 2017). Of other secondary metabolites, Liu et al. (2005) found that condensed tannin concentrations increased when exposed to O 3 ; however, other research has shown that O 3 has no appreciable effect on condensed tannins (Lindroth et al. 2001). These variations in the quantity and quality of litter could affect soil macrofauna, particularly detritivores that feed on litter. ...
Article
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Tropospheric ozone (O3) is a secondary pollutant that causes oxidative stress in plants due to the generation of excess reactive oxygen species (ROS). Phenylpropanoid metabolism is induced as a usual response to stress in plants, and induction of key enzyme activities and accumulation of secondary metabolites occur, upon O3 exposure to provide resistance or tolerance. The phenylpropanoid, isoprenoid, and alkaloid pathways are the major secondary metabolic pathways from which plant defense metabolites emerge. Chronic exposure to O3 significantly accelerates the direction of carbon flows toward secondary metabolic pathways, resulting in a resource shift in favor of the synthesis of secondary products. Furthermore, since different cellular compartments have different levels of ROS sensitivity and metabolite sets, intracellular compartmentation of secondary antioxidative metabolites may play a role in O3-induced ROS detoxification. Plants’ responses to resource partitioning often result in a trade-off between growth and defense under O3 stress. These metabolic adjustments help the plants to cope with the stress as well as for achieving new homeostasis. In this review, we discuss secondary metabolic pathways in response to O3 in plant species including crops, trees, and medicinal plants; and how the presence of this stressor affects their role as ROS scavengers and structural defense. Furthermore, we discussed how O3 affects key physiological traits in plants, foliar chemistry, and volatile emission, which affects plant–plant competition (allelopathy), and plant–insect interactions, along with an emphasis on soil dynamics, which affect the composition of soil communities via changing root exudation, litter decomposition, and other related processes.
... Evaluation of O 3 -induced impacts on nutrient translocation in plants from older to newer leaves has revealed an early shift in the micro-and macronutrients content in the leaf litter, at the outset of the stress exposure, which influences the ecological stoichiometry (Shi et al. 2017). Of other secondary metabolites, Liu et al. (2005) found that condensed tannin concentrations increased when exposed to O 3 ; however, other research has shown that O 3 has no appreciable effect on condensed tannins (Lindroth et al. 2001). These variations in the quantity and quality of litter could affect soil macrofauna, particularly detritivores that feed on litter. ...
... (2) plants will increase their concentrations of leaf nutrients in response to elevated O 3 to increase their resistance to O 3 stress (Cao et al., 2016); and (3) O 3 increases the concentrations of nutrient in woody tissues by inducing the reallocation of leaf nutrients (Samuelson et al., 1996;Shi et al., 2017). However, some studies have also shown that the effects of O 3 on the nutrient concentration of different organs were significantly reduced (e.g., Lindroth et al., 2001) or had no effect (e.g., Li et al., 2019). Differences in leaf longevity, growth habit, species and genotypes, measurement periods, O 3 treatments and environmental conditions may complicate the response of nutrients to O 3. For example, leaf ecological stoichiometry and nutrient re-translocation of the deciduous trees Japanese white birch (Betula platyphylla), Mongolian oak (Q. ...
Article
Ground-level ozone (O3) is a secondary air pollutant and affects the roots and soil processes of trees. Therefore, O3 can affect the uptake and allocation of nutrients in trees, which merits further clarification. A fumigation experiment with five O3 levels was conducted in 15 open top chambers for two poplar clones, and the concentrations of six macronutrients (N, P, K, S, Ca, Mg) in different organs and leaf positions were determined. Under all O3 levels, the concentration of mobile nutrients (N and P) was higher in upper leaves than in lower leaves, while the non-mobile nutrients (Ca and S) concentration was the opposite. Relative to charcoal filtered ambient air (CF), high O3 treatment (NF60) significantly increased the concentration of mobile nutrients K and Mg in upper leaves by 38 % and 33 %, in lower leaves by 142 % and 65 %, respectively, which suggested the effect of O3 on their concentrations was greater at the lower leaf position than at the upper leaf position. Elevated O3 significantly increased the macronutrient concentrations in most organs. The effects of O3 on nutrient concentrations were attributed using graphical vector analysis, suggested that the increase of nutrient concentration in the shoots was attributed to excessive nutrient stocks, while their increase in root was attributed to the “concentration” effect. Compared to CF, NF60 also reduced the root-to-shoot ratio of N, P, S, K, Ca and Mg stocks by 34 %, 39 %, 37 %, 64 %, 46 % and 42 %, respectively, indicating the allocation of increased nutrients to shoots in response to O3 stress. Changes in the allocation pattern of nutrients in different leaf positions and organs of poplar were primarily in response to O3 stress since these nutrients play important roles in some physiological processes. These results will help improve the plantation nutrient utilization by optimizing fertilizer management regimes under O3 pollution.
... (2) plants will increase their concentrations of leaf nutrients in response to elevated O 3 to increase their resistance to O 3 stress (Cao et al., 2016); and (3) O 3 increases the concentrations of nutrient in woody tissues by inducing the reallocation of leaf nutrients (Samuelson et al., 1996;Shi et al., 2017). However, some studies have also shown that the effects of O 3 on the nutrient concentration of different organs were significantly reduced (e.g., Lindroth et al., 2001) or had no effect (e.g., Li et al., 2019). Differences in leaf longevity, growth habit, species and genotypes, measurement periods, O 3 treatments and environmental conditions may complicate the response of nutrients to O 3. For example, leaf ecological stoichiometry and nutrient re-translocation of the deciduous trees Japanese white birch (Betula platyphylla), Mongolian oak (Q. ...
Article
Ground level ozone (O3) pollution is a common environmental stress for plants in many regions of the world, while plant O3 sensitivity is largely influenced by plant water status. Arbuscular mycorrhizal (AM) fungi can form symbiosis with most terrestrial plants and play important role in plant resistance to various environmental stresses. However, how AM symbiosis and water conditions affect plant O3 sensitivity and the underlying mechanism remain unclear. In this study, alfalfa (Medicago sativa L.) was used as test plant, and the effects of AM inoculation and water conditions on plant growth and stress physiology under O3 enrichment were investigated. The results showed that O3 enrichment inhibited plant growth and caused visible injury on plant leaves. Water deficiency also caused significant decrease in plant biomass, while AM symbiosis promoted plant growth regardless of water condition and O3 enrichment. In general, water deficiency decreased while AM symbiosis increased plant O3 sensitivity. Nevertheless, under severe water deficiency, AM inoculation promoted plant growth without increasing plant O3 sensitivity. Water deficiency increased antioxidant enzyme activities and decreased stomatal conductance, while AM symbiosis increased both antioxidant enzyme activities and stomatal conductance. A significant negative correlation between stomatal conductance and O3 effect on plant total biomass supported that both water condition and AM symbiosis affect O3 sensitivity mainly through mediating stomatal conductance. However, AM symbiosis and water condition may differ in regulation mechanisms for stomatal conductance, as AM colonization significantly increased stomatal density, while water deficiency decreased stomatal aperture. Our study suggested that AM inoculation in combination with proper water management could largely mitigate the negative effects of O3 on plants.
... The few reports to date exploring O 3 -related changes in plant nutrient resorption have yielded inconsistent results. In some cases, O 3 was found to reduced NRE ( Gyu et al., 2015 ;Uddling et al., 2005 ), whereas in other cases it enhanced such NRE ( Shang et al., 2018 ;Temple and Riechers, 1995 ) or had no effect ( Baker and Allen, 1995 ;Lindroth et al., 2001 ). These variable findings may be related to the species-specific effects of O 3 or to differences in the O 3 dose and the nutrient availability in the soil ( Shang et al., 2018 ). ...
Article
Concurrent ground-level ozone (O3) pollution and anthropogenic nitrogen (N) deposition can markedly influence dynamics and productivity in forests. Most studies evaluating the functional traits responses of rapid-turnover organs to O3 have specifically examined leaves, despite fine roots are another major source of soil carbon and nutrient input in forest ecosystems. How elevated O3 levels impact fine root biomass and biochemistry remains to be resolved. This study was to assess poplar leaf and fine root biomass and biochemistry responses to five different levels of O3 pollution, while additionally examining whether four levels of soil N supplementation were sufficient to alter the impact of O3 on these two organs. Elevated O3 resulted in a more substantial reduction in fine root biomass than leaf biomass; relative to leaves, more biochemically-resistant components were present within fine root litter, which contained high concentrations of lignin, condensed tannins, and elevated C:N and lignin: N ratios that were associated with slower rates of litter decomposition. In contrast, leaves contained more labile components, including nonstructural carbohydrates and N, as well as a higher N:P ratio. Elevated O3 significantly reduced labile components and increased biochemically-resistant components in leaves, whereas they had minimal impact on fine root biochemistry. This suggests that O3 pollution has the potential to delay leaf litter decomposition and associated nutrient cycling. N addition largely failed to affect the impact of elevated O3 levels on leaves or fine root chemistry, suggesting that soil N supplementation is not a suitable approach to combating the impact of O3 pollution on key functional traits of poplars. These results indicate that the significant differences in the responses of leaves and fine roots to O3 pollution will result in marked changes in the relative belowground roles of these two litter sources within forest ecosystems, and such changes will independently of nitrogen load.
... Genotype by nitrogen enrichment experiments are worthy of attention because several studies have suggested that Populus spp. growth or population dynamics may be sensitive to environmental change factors (Lindroth et al., 2001), including anthropogenic nitrogen enrichment (Kochy and Wilson, 2001;Rogers et al., 2009). ...
Article
Boreal forests receive nitrogen-(N)-enrichment via atmospheric deposition and industrial fertilization. While it is known that N-enrichment can intensify interactions with natural antagonists, it remains poorly understood how genetic variability in plant defense chemistry can affect biotic interactions and height growth in N-enriched environments. We grew replicates of five low- and high-tannin Populus tremula genotypes, respectively, under three N-treatments (ambient, 15, and 150 kg N ha⁻¹ yr⁻¹). We assessed shoot blight occurrence (i.e. symptoms caused by Venturia fungi) during four growing seasons, and tree height growth during the same period. Damage by Venturia spp. increased with N-addition during all years, likely due to enhanced foliar quality. Low–tannin plants showed higher incidences of Venturia infection than high-tannin plants, regardless of the N-input-level. Height responded to an N-by-tannin-group interaction, which occurred because high-tannin plants grew taller than low-tannin plants at the high N-treatment, but not under the other N-levels. This pattern indicates that innate resource investment into tannin production yields a positive effect on growth under N-enriched conditions. Given that N-deposition is increasing globally, our research suggests that further studies are needed to investigate how N-enrichment interacts with plant defense traits globally. Moreover, our research suggests that N-deposition may provide an advantage for well-defended, high-tannin plants; and further, that genetic diversity in plant defense may be a key mechanism by which plant populations respond to this change.
... Carbon flux to and from the soil is also altered by changes in leaf litter quality, altered rhizodeposition of C, changes in soil microbial community composition, and altered soil processes (AINSWORTH, 2017). Tropospheric ozone has the capacity to impact on nutrient cycling by both direct and indirect mechanisms: All of these have the capacity either, independently or in concert, to ultimately reduce the long-term sustainability of ecosystems (LINDROTH et al., 2001, SUN et al., 2012. Tropospheric ozone is known to alter stomatal responses to environmental and in the short term (at higher concentrations) can cause stomata (leaf pores) to close, however, under prolonged chronic exposure (at lower concentrations) many reports document ozone-induced stomatal opening or loss of stomatal sensitivity to closing stimuli, such as drought, light and humidity (MILLS et al., 2013). ...
Article
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One of the most phytotoxic air pollutants is ozone, which can cause considerable damage to vegetation: visible leaf injury, reduction in yield quantity and quality and reduction in photosynthesis, alterations to carbon allocation, and in the sensitivity to biotic and abiotic stresses. Ozone is a secondary pollutant and prevailed at high concentrations over rural regions. Moreover, ozone concentrations are expected to increase in the future and will continue to be a serious threat to crop productivity. This paper presents a comprehensive review the undertaken studies of ozone's impacts on crops and natural ecosystems.
... Insect herbivores are expected to damage plants more if O 3 decreases the rate of C assimilation so that less C is available for C-based defensive chemicals (74). Numerous studies have demonstrated that O 3 affects phenolics and terpenes, especially in angiosperms (75), although different groups of phenolics and terpenes may respond differently to O 3 (19,76,77). Despite the recent advancements in the understanding of O 3 effects on foliar quality, O 3 effects on latex and other constituents of saps exuded from damaged tissues remain completely unknown, although plant latex plays an important role in defense against herbivores (78). ...
Article
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Elevated tropospheric ozone concentrations induce adverse effects in plants. We reviewed how ozone affects (i) the composition and diversity of plant communities by affecting key physiological traits; (ii) foliar chemistry and the emission of volatiles, thereby affecting plant-plant competition, plant-insect interactions, and the composition of insect communities; and (iii) plant-soil-microbe interactions and the composition of soil communities by disrupting plant litterfall and altering root exudation, soil enzymatic activities, decomposition, and nutrient cycling. The community composition of soil microbes is consequently changed, and alpha diversity is often reduced. The effects depend on the environment and vary across space and time. We suggest that Atlantic islands in the Northern Hemisphere, the Mediterranean Basin, equatorial Africa, Ethiopia, the Indian coastline, the Himalayan region, southern Asia, and Japan have high endemic richness at high ozone risk by 2100. This is an open-access publication and can be downloaded at now cost: https://advances.sciencemag.org/content/6/33/eabc1176
... Although some previous meta-analyses have investigated the responses of N:P ratios to global changes at the whole-plant level 10 , the responses at the foliar level have received less attention. In addition, the responses of foliar [C] and foliar C:N ratio to global change have received less attention [11][12][13] , and there is very little information about changes in foliar stoichiometry in response to warming and increased precipitation [14][15][16] . Therefore, it is urgent to investigate the responses of foliar C:N and N:P ratios to global change, including warming, changes in precipitation, N deposition and eCO 2 . ...
Article
Full-text available
Foliar-level stoichiometry plays an important role in ecosystem elemental cycling. Shifts in foliar ratios of carbon to nitrogen (C:N) and nitrogen to phosphorus (N:P) in response to global change can therefore have a large impact upon ecosystem function. We conducted a meta-analysis with 2,236 paired observations from 123 published studies to investigate the responses of foliar C:N and N:P ratios to experimental global change treatments, i.e. warming, increased precipitation, drought, N addition and elevated carbon dioxide concentration (eCO2), in field conditions. Foliar C:N and N:P ratios were neither affected by warming nor by increased precipitation. Foliar C:N ratio increased with drought and eCO2, and decreased with N addition. Foliar N:P ratios declined with eCO2, and increased under drought and N addition. Our results suggested the responses of the C:N ratio to global change were mainly related to shifts in foliar [N], whereas changes in the N:P ratio were related to the responses of both [N] and [P]. Moreover, the response magnitude of foliar N:P ratio decreased with treatment duration under increased precipitation, N addition and eCO2. Our findings are important for our understanding of plant nutrient dynamic and modeling of nutrient biogeochemistry under global change.
... Elevated CO 2 also stimulate the soil microbial activity under various RCTs compared to control through increased concentration of non-structural carbohydrates in leaves. Consequently leaf litters with higher concentration of carbohydrates and amino acids and lower concentration of phenolic acids provides more labile carbon to microbes (Lindroth et al. 2001). ...
... This leads to depressed root growth and the potential for affected species to have an increased susceptibility to drought. Ozone and accompanying N deposition alters elemental ratios in foliage, litter, soil (Boerner and Rebbeck 1995;Anderson et al. 2001;Lindroth et al. 2001) and global ecosystems ( Peñuelas et al. 2013). Concentrations of tannins, lignin, and phenolics ( Kim et al. 1998;Findlay et al. 1996;(Baumgarten et al. 2000;Saleem et al. 2001) are also affected by O 3 exposure. ...
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Tropospheric ozone (O3) is an important stressor in natural ecosystems, with well‐documented impacts on soils, biota, and ecological processes. The effects of O3 on individual plants and processes scale up through the ecosystem through effects on carbon, nutrient, and hydrologic dynamics. Ozone effects on individual species and their associated microflora and fauna cascade through the ecosystem to the landscape level. Systematic injury surveys demonstrate that foliar injury occurs on sensitive species throughout the globe. However, deleterious impacts on plant carbon, water, and nutrient balance can also occur without visible injury. Because sensitivity to O3 may follow coarse physiognomic plant classes (in general, herbaceous crops are more sensitive than deciduous woody plants, grasses, and conifers), the task still remains to use stomatal O3 uptake to assess class and species’ sensitivity. Investigations of the radial growth of mature trees, in combination with data from many controlled studies with seedlings, suggest that ambient O3 reduces growth of mature trees in some locations. Models based on tree physiology and forest stand dynamics suggest that modest effects of O3 on growth may accumulate over time, other stresses (prolonged drought, excess nitrogen deposition) may exacerbate the direct effects of O3 on tree growth, and competitive interactions among species may be altered. Ozone exposure over decades may be altering the species composition of forests currently, and as fossil fuel combustion products generate more O3 than deteriorates in the atmosphere, into the future as well. This article is protected by copyright. All rights reserved.
... There have indeed been many studies on the effects of O 3 on leaf N concentration, but their results are inconsistent with each other. Elevated O 3 increased leaf N concentration (e.g., Cao et al., 2016;Shang et al., 2018), had no significant effects (e.g., Häikiö et al., 2006;Oksanen et al., 2005), or reduced it (e.g., Lindroth et al., 2001;Singh et al., 2009). The past studies are also inconsistent in their reported units of the leaf N concentration, which was expressed on leaf mass basis in some studies (e.g., Oksanen et al., 2005;Shang et al., 2018), but was based on per unit leaf area in other studies (e.g., Kitao et al., 2009;Oikawa and Ainsworth, 2016). ...
Article
We investigated the effects of elevated ozone (O3) concentration on leaf nitrogen (N), a key determinant of plant photosynthesis, with two clones of poplar grown in open-top chambers. We focus on the difference between mass-based leaf N concentration (Nmass) and area-based one (Narea) in their responses to elevated O3, and the allocation of N to different leaf components: photosynthetic apparatus, cell walls, and others under elevated O3 level. Our results showed that elevated O3 significantly increased Nmass, but reduced Narea and leaf mass per area (LMA). The two clones showed no difference in Nmass response to O3, but the more sensitive clone showed greater reduction of Narea and LMA due to O3. We also found positive relationships between Narea and photosynthetic parameters, e.g. light-saturated photosynthetic rate (Asat). Furthermore, elevated O3 significantly reduced photosynthetic N-use efficiency (PNUE) and leaf N allocation to photosynthetic components, while increasing N allocation to cell walls and other components. We concluded that plants invested more N in cell walls and other components to resist O3 damages at the expense of photosynthetic N. The change of N allocation in plant leaves in response to elevated O3 could have an impact on ecological processes, e.g. leaf litter decomposition.
... This variation is tightly related to the position of the specie in the crown [9], nitrogen fertility level, season and co-occurring pollutant concentrations [10]. Nitrogen level decreases in the foliage of trees growing under elevated atmospheric CO 2 [11][12][13]. It is also decreased in the litter [14]. ...
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Despite the increase of the atmospheric CO2, cork oak (Quercus suber L.) forests don’t stop degenerating. Deficits in water balance and in nutritional elements might be the main raisons. Standing as a potential regulator of the ecosystem nutrient dynamics, leguminous plants (Fabaceae, Leguminosae) are a good test case for individual species effects on Tunisian forests. They are the most diverse and widespread group of plants with the capacity of N2 fixation, and are particularly abundant in Kroumiri forests. The use of natural legume species in Kroumirie, as soil fertility and grassland productivity enhancer, could be very interesting in the assessment of a new national land exploitation strategy aiming to increase carbon sequestration and climate change mitigation. Understanding the impact of these legume species on cork oak, using air nitrogen-fixation technique computation, would be crucial for future land management and policy decisions. A morphological and eco-physiological study of the Cytisus triflorus plant associated with Cork oak was carried out in Tabbouba, at Nefza region. Three sites were selected: CM1 (Cytisus triflorus alone), CM2 (Cork oak associated with Cytisus triflorus) and CM3 (Cork oak only). Morphological (height, diameter, density), physiological (stomatal conductance, water potential, transpiration, photosynthesis) and hydric parameters were measured for the two species. The morphological study results showed no significant difference sites for each species except for density parameter. On the other hand, physiological parameters measured for oaks trees clearly manifested significant differences in photosynthesis, transpiration and hydric conductance between CM2 and CM3 sites. The cork oak in association with the Cytisus are in better growth and productivity conditions than when they are alone.
... Several studies have shown that the elevated CO 2 increased lepidopteran insect feeding and damage severity in agricultural crops Lindroth et al., 2001), because of the increased proportion of C:N in host plant tissue and lower nutritional quality caused by elevated CO 2 ( . For example, larvae of Helicoverpa armigera fed on wheat grown in elevated CO 2 showed the extended larval life span and increased consumption with reduced growth rate (Chen, Wu & Ge, 2004). ...
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Background Bt crops will face a new ecological risk of reduced effectiveness against target-insect pests owing to the general decrease in exogenous-toxin content in Bt crops grown under elevated carbon dioxide (CO 2 ). The method chosen to deal with this issue may affect the sustainability of transgenic crops as an effective pest management tool, especially under future atmospheric CO 2 level raising. Methods In this study, rhizobacterias, as being one potential biological regulator to enhance nitrogen utilization efficiency of crops, was selected and the effects of Bt maize (Line IE09S034 with Cry1Ie vs. its parental line of non- Bt maize Xianyu 335) infected by Azospirillum brasilense (AB) and Azotobacter chroococcum (AC) on the development and food utilization of the target Mythimna separate under ambient and double-ambient CO 2 in open-top chambers from 2016 to 2017. Results The results indicated that rhizobacteria infection significantly increased the larval life-span, pupal duration, relative consumption rate and approximate digestibility of M. separata , and significantly decreased the pupation rate, pupal weight, adult longevity, fecundity, relative growth rate, efficiency of conversion of digested food and efficiency of conversion of ingested food of M. separata fed on Bt maize, while here were opposite trends in development and food utilization of M. separata fed on non- Bt maize infected with AB and AC compared with the control buffer in 2016 and 2017 regardless of CO 2 level. Discussion Simultaneously, elevated CO 2 and Bt maize both had negative influence on the development and food utilization of M. separata . Presumably, CO 2 concentration arising in future significantly can increase their intake of food and harm to maize crop; however, Bt maize infected with rhizobacterias can reduce the field hazards from M. separata and the application of rhizobacteria infection can enhance the resistance of Bt maize against target lepidoptera pests especially under elevated CO 2 .
... Therefore, if the growth of Douglas-fir trees in our site had been limited by nutrient depletion through biomass harvesting, then the foliar carbon concentration should have been lower in the greatest removal level (removal of all woody biomass down to a 2.5 cm top). Although the responses of foliar carbon can vary with many factors [66,67], the result of our study-on our cool and moist study site there was no difference in foliar carbon-may be consistent with the result of the foliar N contents. ...
Article
To investigate the long-term impacts of biomass harvesting on site productivity, we remeasured trees in the 1974 Forest Residues Utilization Research and Development Program at Coram Experimental Forest in western Montana. Three levels (high, medium, and low) of biomass removal intensity combined with broadcast burning treatment were assigned after clearcut in western larch (Larix occidentalis Nutt.) stands in 1974. From 1976 to 79, twenty five 2 + 0 bare root seedlings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were consecutively planted in rows. In 2013, tree height, dbh (diameter at breast height), foliar N and C concentrations were measured. From cross-sectional sapwood area, growth efficiency (the ratio of 5-year-basal area increment to total leaf area) was calculated. Previous measurements from 1980, 1987, 1992, and 2001 were used for dbh and height growth analyses. At this site, none of the response variables were affected by biomass removal level. Only seedling planting year contributed significantly to affect tree mean height, dbh, volume. Growth efficiency was not affected by any treatment. These results indicate no apparent effect of biomass removal on site productivity for the range of biomass harvest levels performed.
... Depending on the cell injury, nutrient translocation from older to younger leaves is changed by elevated ozone, so ozone can affect the nutrient resorption of mobile nutrients. There are some studies on the effects of ozone on nutrient resorption, but their results are inconsistent (Gyu et al., 2015;Lindroth et al., 2001;Temple and Riechers, 1995). Therefore, this study can provide evidence for the effect of ozone on nutrient resorption, and it is necessary to better understand the impact of ozone on plant nutrient use. ...
Article
The effects of elevated ozone on C (carbon), N (nitrogen) and P (phosphorus) ecological stoichiometry and nutrient resorption in different organs including leaves, stems and roots were investigated in poplar clones 546 (P. deltoides cv. '55/56' × P. deltoides cv. 'Imperial') and 107 (P. euramericana cv. '74/76') with a different sensitivity to ozone. Plants were exposed to two ozone treatments, NF (non-filtered ambient air) and NF60 (NF with targeted ozone addition of 60 ppb), for 96 days in open top chambers (OTCs). Significant ozone effects on most variables of C, N and P ecological stoichiometry were found except for the C concentration and the N/P in different organs. Elevated ozone increased both N and P concentrations of individual organs while for C/N and C/P ratios a reduction was observed. On these variables, ozone had a greater effect for clone 546 than for clone 107. N concentrations of different leaf positions ranked in the order upper > middle > lower, showing that N was transferred from the lower senescent leaves to the upper ones. This was also indicative of N resorption processes, which increased under elevated ozone. N resorption of clone 546 was 4 times larger than that of clone 107 under ambient air (NF). However, elevated ozone (NF60) had no significant effect on P resorption for both poplar clones, suggesting that their growth was only limited by N, while available P in the soil was enough to sustain growth. Understanding ecological stoichiometric responses under ozone stress is crucial to predict future effects on ecological processes and biogeochemical cycles.
... The increased proportion of C: N in plant tissues reduced its nutritional quality which induced increased feeding by insect herbivores for sufficient nutritional intake, thus greater amount of herbivory and crop damage (Stitt & Krapp, 1999). Many studies have shown that Lepidoptera insects increase consumption (food intake), and the damage is more severe when the crops are grown under elevated CO 2 concentration (Lindroth et al., 2001). For example, larvae of Helicoverpa armigera fed on wheat grown in elevated CO 2 showed the extended larval life-span and increased consumption with reduced growth rate (Chen, Wu, & Ge, 2004). ...
Article
Effects of elevated atmospheric CO2 (elevated CO2 vs. ambient CO2) and temperature (+0.67–0.79°C vs. ambient temperature) on the developmental life cycle of Spodoptera litura and the food utilization of the fourth-instar larvae fed on soybean (resistant cultivar Lamar vs. susceptible landrace JLNMH) grown in open-top chambers were studied from 2013 to 2015. The results indicated that: (i) compared with ambient CO2, elevated CO2 significantly prolonged the duration of larva and pupa, and adult longevity; significantly decreased the pupation rate, pupal weight, fecundity, the relative growth rate (RGR), efficiency of conversion of ingested food (ECI) and efficiency of conversion of digested food (ECD); and increased the relative consumption rate (RCR) and approximate digestibility (AD). (ii) Compared with ambient temperature, elevated temperature significantly shortened the duration of larva and pupa; significantly decreased the pupal weight; and increased the RGR, RCR, ECD and ECI. (iii) Compared with the susceptible soybean accession JLNMH, the resistant soybean cultivar Lamar significantly prolonged the duration of larva and pupa; significantly decreased the pupation rate, pupal weight, adult longevity, fecundity and RGR, RCR and AD; and increased the indexes of ECD. (iv) At elevated temperature, S. litura fed on resistant vs. susceptible cultivars showed opposite trends in the RGR, RCR, AD, ECD and ECI. In addition, elevated temperature under elevated CO2 significantly decreased the RGR (2014), ECD (2013 & 2014) and ECI (2013) and increased the AD (2013 & 2014) compared with other treatment combinations when S. litura fed on Lamar. Future climatic change of temperature and CO2 concentration would likely affect growth and food utilization of S. litura, with increased food intake, but the reduced fecundity may compensate for the increased food consumption, resulting in no significant reduction in insect-induced yield loss in soybean production. Nevertheless, use of insect resistant soybean cultivars will aid in ecological management of S. litura and reduce the insecticide load in soybean production.
... Decomposition is often positively related to residue N content(Hobbie et al., 2012), but contrasting results have been reported for the impact of O 3 on litter N concentration. WhereasKing, Liu, and Aspinwall (2013) reported that elevated O 3 causes a general decrease in litter N concentration, others have found an increase(Lindroth et al., 2001), which may explain why litter decomposition differs between species(Williamson, Mills, & Freeman, 2010). Decomposition of SOM by fungi decreases in elevated O 3(Edwards & Zak, 2011;Yue et al., 2015) more than decomposition by bacteria(Zhang et al., 2014), suggesting that any effect of O 3 on decomposition could be lower in productive systems with soil communities dominated by highly active bacteria than in systems with lower productivity where fungi and less active bacteria dominate. ...
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Risks associated with exposure of individual plant species to ozone (O3) are well documented, but implications for terrestrial biodiversity and ecosystem processes have received insufficient attention. This is an important gap because feedbacks to the atmosphere may change as future O3 levels increase or decrease, depending on air quality and climate policies. Global simulation of O3 using the Community Earth System Model (CESM) revealed that in 2000, about 40% of the Global 200 terrestrial ecoregions (ER) were exposed to O3 above thresholds for ecological risks, with highest exposures in North America and Southern Europe, where there is field evidence of adverse effects of O3, and in central Asia. Experimental studies show that O3 can adversely affect the growth and flowering of plants and alter species composition and richness, although some communities can be resilient. Additional effects include changes in water flux regulation, pollination efficiency, and plant pathogen development. Recent research is unraveling a range of effects belowground, including changes in soil invertebrates, plant litter quantity and quality, decomposition, and nutrient cycling and carbon pools. Changes are likely slow and may take decades to become detectable. CESM simulations for 2050 show that O3 exposure under emission scenario RCP8.5 increases in all major biomes and that policies represented in scenario RCP4.5 do not lead to a general reduction in O3 risks; rather, 50% of ERs still show an increase in exposure. Although a conceptual model is lacking to extrapolate documented effects to ERs with limited or no local information, and there is uncertainty about interactions with nitrogen input and climate change, the analysis suggests that in many ERs, O3 risks will persist for biodiversity at different trophic levels, and for a range of ecosystem processes and feedbacks, which deserves more attention when assessing ecological implications of future atmospheric pollution and climate change.
... Therefore, elevated CO 2 concentrations have far-reaching effects on photosynthesis and yield (Agrell et al. 2000). Such concentrations cause the greenhouse effect, which can gradually warm the climate, influence water balance, alter seasonal precipitation patterns, and vary water resources regionally (Richard et al. 2001;Duan et al. 2013). Moreover, elevated temperature increases potential evaporation, causing the most arid regions on earth to become more arid (Rind et al. 1990). ...
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The variation of net photosynthetic rate (Pn), transpiration rate (Tr), and intercellular CO2 concentration (Ci) with changes in light intensity of Tamarix ramosissima leaves was studied during the growing seasons of 2012 and 2013 in the extremely arid region of Ejina Oasis, using the LI-6400 portable photosynthesis measurement system. Results indicate that Ci decreased and Pn and Tr increased with light intensity. After reaching the light saturation point, Pn decreased gradually with the enhanced greater light intensity. The apparent maximum photosynthetic rate increased with CO2 concentration, from 17.69 to 27.05 µmol/(m² s); apparent quantum efficiency also increased, as did the light saturation point of T. ramosissima leaves. Water use efficiency (WUE) increased initially but gradually decreased after maximizing with the enhanced light intensity. Tr and Ci increased with elevated CO2 concentration. Pn and WUE increased with CO2 concentration when that concentration was 200–600 µmol/mol. When that concentration increased to 1000 µmol/mol, WUE increased initially but eventually decreased.
... Effects of O3 on (a) relative N fertilizer efficiency in wheat and (b) the resorption of leaf N in trees before litterfall, where ECLAIRE results are compared with previous studies. Species are silver birch (solardomes;Uddling et al., 2006) and paper birch(Lindroth et al., 2001) (WP9 activity). ...
... For example, tannins prevent stress through protection from photodamage (Close & McArthur 2002) and alter nutrient cycling by reducing litter decomposition rates, forming protein complexes , microbial priming, or directly inhibiting microbial functioning (H€ attenschwiler & Vitousek 2000; Kraus, Dahlgren & Zasoski 2003; Kraus et al. 2004; Madritch, Jordan & Lindroth 2007; Schweitzer et al. 2008; Madritch & Lindroth 2015 ). Foliar concentrations of condensed tannins vary considerably among tree species and genotypes, and in both aspen and paper birch are influenced by environmental variation (Lindroth et al. 2001; Osier & Lindroth 2001; Donaldson & Lindroth 2007). Tannins also comprise a significant carbon fraction in plants, behind only structural compounds (Hernes & Hedges 2000), and comprise upwards of 35% of foliar dry mass in aspen and paper birch. ...
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Spectroscopy has recently emerged as an effective method to accurately characterize leaf biochemistry in living tissue through the application of chemometric approaches to foliar optical data, but this approach has not been widely used for plant secondary metabolites. Here, we examine the ability of reflectance spectroscopy to quantify specific phenolic compounds in trembling aspen ( Populus tremuloides ) and paper birch ( Betula papyrifera ) that play influential roles in ecosystem functioning related to trophic‐level interactions and nutrient cycling. Spectral measurements on live aspen and birch leaves were collected, after which concentrations of condensed tannins (aspen and birch) and salicinoids (aspen only) were determined using standard analytical approaches in the laboratory. Predictive models were then constructed using jackknifed, partial least squares regression ( PLSR ). Model performance was evaluated using coefficient of determination ( R ² ), root‐mean‐square error ( RMSE ) and the per cent RMSE of the data range (% RMSE ). Condensed tannins of aspen and birch were well predicted from both combined ( R ² = 0·86, RMSE = 2·4, % RMSE = 7%)‐ and individual‐species models (aspen: R ² = 0·86, RMSE = 2·4, % RMSE = 6%; birch: R ² = 0·81, RMSE = 1·9, % RMSE = 10%). Aspen total salicinoids were better predicted than individual salicinoids (total: R ² = 0·76, RMSE = 2·4, % RMSE = 8%; salicortin: R ² = 0·57, RMSE = 1·9, % RMSE = 11%; tremulacin: R ² = 0·72, RMSE = 1·1, % RMSE = 11%), and spectra collected from dry leaves produced better models for both aspen tannins ( R ² = 0·92, RMSE = 1·7, % RMSE = 5%) and salicinoids ( R ² = 0·84, RMSE = 1·4, % RMSE = 5%) compared with spectra from fresh leaves. The decline in prediction performance from total to individual salicinoids and from dry to fresh measurements was marginal, however, given the increase in detailed salicinoid information acquired and the time saved by avoiding drying and grinding leaf samples. Reflectance spectroscopy can successfully characterize specific secondary metabolites in living plant tissue and provide detailed information on individual compounds within a constituent group. The ability to simultaneously measure multiple plant traits is a powerful attribute of reflectance spectroscopy because of its potential for in situ – in vivo field deployment using portable spectrometers. The suite of traits currently estimable, however, needs to expand to include specific secondary metabolites that play influential roles in ecosystem functioning if we are to advance the integration of chemical, landscape and ecosystem ecology.
... L'absence d'effet significatif du doublement des concentrations actuelles de CO 2 sur la masse racinaire (Fig. 3) ainsi que sur la fréquence des différentes phases de développement (Fig. 4) est en accord avec la conclusion de plusieurs auteurs [27,29,43] selon lesquels le premier effet de l'augmentation du niveau de CO 2 sur la croissance se fait via l'accumulation du carbone issu de la photosynthèse sous forme d'amidon dans les feuilles. Dans notre étude, l'augmentation de la photosynthèse des plants de haricots croissant sous un régime riche en CO 2 jusqu'au 66 e jour après semis s'accompagne d'une augmentation significative du contenu en sucres dans les feuilles. ...
Article
The effects of elevated CO2 on bean (Phaseolus vulgaris) physiology were analysed in greenhouses at 350 μl/l and at 700 μl/l (i.e. twice the actual level). The photosynthesis of beans grown under high CO2 concentration increased (+73% until the 40th day after germination). At high concentration, leaf nitrogen content was 15% lower while the C/N ratio was 19% higher. Leaf soluble and unsoluble sugar contents increased (+49 and +64%, respectively). The CO2 doubling significantly increased leaf area by 36% and leaf mass by 57%. Two months after sowing, plants exposed to 700 μl/1 produced pods with higher fresh and dry weights (+44 and +110%, respectively). The individual pod dry weight and length also were also higher (+100 and +10%) but the number of seeds per pod was reduced (-9%). The soluble sugar content increased by 41%. In summary, the results suggest that the CO2 doubling has an important effect on bean yield, particularly on the quantitative level, but the increase in the soluble sugar content was more important on the qualitative level.
... However, there were no effects of eCO 2 on leaf N resorption efficiency in past studies in FACE experiments (e.g. Finzi et al., 2001;Lindroth et al., 2001;Norby & Iversen, 2006;Esmeijer-Liu et al., 2009) or in a review of earlier experiments (Norby et al., 2001). Mini-FACE experiments (1-4 m in diameter) were excluded from the present study out of concern for potential large edge effects associated with the small CO 2 enrichment footprint. ...
... Therefore, any significant alteration in its levels is likely to cause a marked effect on plant metabolism and growth (Pattanayak and Tripathy 2011;Biswal et al. 2012). Elevated [CO 2 ] tends to alter the foliar chemistry of plants (Lindroth et al. 2001). The Chl and leaf protein contents were lower by 3-5 and 1.5-3 %, respectively, in B. juncea plants grown in CO 2 -enriched environment than the ambient ones. ...
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Increased atmospheric [CO2] is likely to affect photosynthesis, plant growth, and yield potential of plants. Mustard (Brassica juncea L.) is an important oil seed crop that is widely grown in India. Therefore, the impact of elevated [CO2] (585 μmol mol−1) on pigment and protein content, chlorophyll a fluorescence, photosynthetic electron transport reactions, CO2 assimilation, biomass production, and seed yield potential was measured in B. juncea cv Pusa Bold, grown inside free air carbon dioxide enrichment (FACE) rings installed on the campus of Jawaharlal Nehru University, New Delhi, India. Plants were grown for three consecutive winter seasons (2010—2013), in ambient (385 μmol mol−1) or elevated [CO2], in field conditions. Elevated [CO2] had no significant effect on the minimal chlorophyll fluorescence (F 0), while the quantum efficiency of Photosystem II, measured as variable fluorescence (F v = F m–F 0) to maximum fluoresence (F m), increased by 3 %. Electron transport rate, photosystem I, photosystem II, and whole chain electron transport rates increased by 8 % in elevated [CO2]. However, the net photosynthesis rate increased by ≈50 % in three growing seasons under elevated [CO2] condition. The stomatal conductance and transpiration rate decreased resulting in higher photosynthetic water use efficiency. The photosynthesizing surface, i.e., leaf area index substantially increased leading to higher biomass and seed yield under elevated [CO2] condition. Acclimatory downregulation of photosynthesis and plant productivity was not observed in three consecutive growing years suggesting that in the absence of nutrient limitation, B. juncea is highly responsive to elevated CO2 whose yield potential shall increase in changing climatic conditions.
... However, prolonged (6-year) droughts would eventually cause severe dieback (Grant et al. 2006). Therefore, some researchers stress that the long-term effects of elevated atmospheric CO 2 on aspen will be complex and difficult to predict (Hogg 2001, Lindroth et al. 2001). ...
... Atmospheric chemical composition affects foliar chemical composition. In general, leaf N concentration of plants is decreased under elevated CO 2 (Koricheva et al. 1998), marginally reduced under elevated 03 (Lindroth et al. 2000), which disagreed with the results of Scherzel et al. (1998), who showed that single pollutants, such as 03, didn't significantly alter foliar N concentration. When treated in combination with CO2, however, leaf N concentration was decreased by 18% to 40% (Scherzel et al. 1998). ...
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In order to evaluate the dynamics of carbon storage during forest succession and explore the significance of water relations and soil stability in forest environments, a study was conducted in 2011. This study investigated the dynamics of soil organic carbon (SOC) fractions and its protection through aggregation along the successional forests. An experiment in South China examined pine forest (PF), pine and broadleaved mixed forest (PBMF), and monsoon evergreen broadleaf forest (MEBF), which represent the early, middle, and advanced succession stages, respectively. Soil was sampled at 0-15, 15-30, 30-45, and 45-60 cm depths. We analyzed active, slow, and passive organic carbon, as well as particulate organic matter carbon (POM-C) and nitrogen (POM-N), and measured the weight and concentration of water-stable aggregates in four classes (>2000 μm, 250-2000 μm, 53-250 μm, and <53 μm). The results suggested that various carbon fractions and the percentage of active and passive carbon to total organic carbon (TOC) increased with forest succession. The percentage of water-stable aggregates in >2000 μm (0-15 cm and 15-30 cm) and <53 μm (45-60 cm) in MEBF was significantly higher than in PBMF and PF. The SOC content of all size classes of water-stable aggregates in 0-45 cm were significantly increased with forest succession. In conclusion, forest succession contributed to the accumulation of carbon storage, and the increasing percentage of silt- and clay-size (<53 μm) fraction improved the stability of soil organic matter (SOM).
Chapter
The expansion of urban areas, the acceleration of traffic, the acceleration of economic growth, and the excessive use of energy are all characteristics of industrialized nations that have contributed to the worsening of air pollution. The integrity of the natural world is compromised by all these elements, which have a domino effect on one another and work together to harm it. A major ecological problem is the regional effects of air pollution on various plant species. Unlike animal populations, plant populations are constantly (24/7) and directly exposed to the danger of pollution. Biochemical, physiological, morphological, and anatomical reactions are among the many ways in which these organisms take in, store, and process contaminants that land on their surfaces. This research aims to find out how two possible therapeutic plant species Catharanthus roseus L. and Ocimum sanctum L. react to different levels of air pollution (vehicular pollution) in terms of their morphology, physiology, biochemistry, and pharmacognosy.
Article
Increased surface ozone (O3) pollution seriously threatens crop production, and ethylenediurea (EDU) can alleviate crop yield reduction caused by O3. However, the reason for the decrease in grain nitrogen (N) accumulation caused by O3 and whether EDU serves as N fertilizer remain unclear. An experiment was conducted to investigate the impacts of factorial combinations of O3 enrichment (ambient air plus 60 ppb) and EDU (foliage spray with 450 ppm solutions) on N concentration, accumulation and remobilization in hybrid rice seedlings. Compared to ambient condition, elevated O3 significantly inhibited the N accumulation in vegetative organs during anthesis and grain N accumulation during the maturity stage. Elevated O3 significantly decreased the total N accumulation duringanthesis and maturity stages, with a greater impact at the latter stage. The decrease in grain N accumulation caused by O3 was attributed to a decrease in N remobilization of vegetative organs during the grain filling period as well as to a decrease in post-anthesis N uptake. However, there was no significant change in the proportion of N remobilization and N uptake in grain N accumulation. The inhibitory effect of O3 on N remobilization in the upper canopy leaves was greater than that in the lower canopy leaves. In addition, elevated O3 increased the N accumulation of panicles at the anthesis stage, mainly by resulting in earlier heading of rice. EDU only increased N accumulation at the maturity stage, which was mainly attributed to an increase in rice biomass by EDU. EDU had no significant effect on N concentration, N remobilization process, and N harvest index. The findings are helpful to better understand the utilization of N fertilizer by rice under O3 pollution, andcan also provide a theoretical basis for sustainable nutrient management to alleviate the negative impact of O3 on crop yield and quality.
Article
Carbon (C), nitrogen (N) and phosphorus (P) concentrations and stoichiometry play important roles in biogeochemical cycles of the ecosystems, yet it is still unclear how the allocations of C, N and P concentrations and stoichiometry among plant organs and soils related to O3 stress and straw return. Here, a pot experiment was conducted in open top chambers to monitor the response of C, N and P concentrations and stoichiometry of leaves, stems, roots and soils during a growing season (branching, flowering and podding stages) of soybean (Glycine max; a species highly sensitive to O3) to background O3 concentration (44.8 ± 5.6 ppb), O3 stress (79.7 ± 5.4 ppb) and straw treatment (no straw return and straw return). O3 stress significantly decreased root biomass. Straw return significantly increased root biomass under O3 stress at branching and flowering stages. Generally, O3 stress and straw return showed significant effects on the C, N and P concentrations of leaves and soils, and stoichiometric ratios of leaves, stems and microbial biomass. The C, N and P concentrations and stoichiometry of leaves, stems, roots and soils in response to O3 stress and straw return at the branching stage were inconsistent with the changes observed at the flowering and podding stages. The P conversion efficiency showed significant relationship with root P concentration under the combined effects of O3 stress and straw return. Altogether, the present study indicated that C, N and P concentrations of soybean might be more important than stoichiometric ratios as a driver of root defence against O3 stress in the case of straw return.
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Elevated atmospheric CO2 concentration (eCO2) typically stimulates tree growth, which is mediated by nitrogen (N) availability; but how N regulates tree biomass responses to eCO2 remains uncertain, which limits our prediction of forest carbon (C) cycling under future global change scenarios. A meta‐analysis of a global dataset including 3,399 observations from 283 papers published from 1980s to February 2021 was conducted with the aim of elucidating N‐mediated responses of tree biomass production to eCO2 and the underlying mechanisms. We found that eCO2 stimulated tree biomass production (+32.0%), while it induced accumulation of non‐structural carbohydrates in leaves rather than in woods and roots, suggesting that the production may be C‐limited but depend on the sink strength of organs. Biomass responses to eCO2 of N‐fertilized trees (+39.6%) were 68.4% greater than those of non‐fertilized trees (+23.5%), confirming that tree growth is also N‐limited. Such N limitation was alleviated by the eCO2‐induced increases in N uptake and N‐use efficiency (NUE), with the former being more important. Increases in tree N pool arose from the enhanced production of fine roots with a lower specific root length, whereas increases in NUE resulted from the flexibility in tissue C:N ratios instead of N resorption efficiency. The positive responses of tree biomass production to eCO2 were greater for ectomycorrhizal trees and conifers than for arbuscular mycorrhizal trees and angiosperms, respectively. Synthesis. Our findings suggest that eCO2 stimulates tree biomass production by increasing C availability, and alleviating N limitation in a feedback way via enhancing N uptake and NUE, and they improve our mechanistic understanding of responses of forest productivity and C sequestration to eCO2 under global change.
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AimsC, N and P ecological stoichiometry plays important roles on biogeochemical cycles in ecosystems, yet the relationship between plant and soil stoichiometry and stoichiometric effects on the growth of soybean root in response to the O 3 stress and straw return remain poorly understood. Methods Here, a pot experiment was conducted in open top chambers to monitor the response of C, N and P ecological stoichiometry of leaves, shoots, roots and soils during a growing season (branching, flowering and podding stages) of soybean (Glycine max; a highly sensitive species to O 3 ) to background O 3 concentration (45 ± 10 ppb), O 3 stress (80 ±10 ppb) and straw treatment (no straw return and straw return). ResultsThe O 3 stress significantly decreased root biomass. The straw return significantly increased root biomass under the O 3 stress at branching and flowering stages. Generally, the O 3 stress and straw return showed significant effects on the C, N, P concentrations of leaves and soils, and stoichiometric ratios of leaves, shoots and microbial biomass. C, N, P concentrations and stoichiometric ratios of leaves, shoots, roots and soils responses to the O 3 stress and straw return at branching stage were inconsistent with changes observed at the flowering and podding stages. The P conversion efficiency showed significant relationship with root P concentration under the combined effects of O 3 stress and straw return. ConclusionsC, N, and P concentrations of soybean might be more important than stoichiometric ratios as a driver of defense against the O 3 stress in the case of straw return.
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Condensed tannins (CTs), synthesized via the phenylpropanoid and malonic acid pathways, occur in Populus tissues at widely varying concentrations. Both concentration and molecular structure are determined by the independent and interactive effects of genetic, ontogenetic, and environmental factors that influence molecular control of CT synthesis, other secondary metabolite production, and plant growth. CTs have a limited role in defending Populus against herbivores, but are associated with pathogen resistance, structuring of herbivore and soil microbial communities, and regulation of soil ecosystem processes (e.g. respiration, decomposition, nutrient cycling) with feedbacks to plant fitness. Shifts in CT expression and distribution resulting from human‐mediated environmental changes are likely to alter organismal interactions, community and ecosystem functions, and evolutionary processes. Future research should aim to strengthen understanding of causal connections between CTs and their biological effects across scales of organization, space and time, and to elucidate how environmental change influences CT production, biochemical tradeoffs, and interrelationships with plant fitness that drive evolutionary processes.
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Retention of carbon (C), either by physical mechanisms or microbial uptake, is a key driver of the transformation and storage of C and nutrients within ecosystems. Both the molecular composition and nutrient content of organic matter influence the rate at which it is retained in streams, but the relative influence of these characteristics remains unclear. We estimated the effects of nutrient content and molecular composition of dissolved organic C (DOC) on uptake in boreal streams by measuring rates of C retention, in situ, following introduction of leachates derived from alder, poplar, and spruce trees subject to long-term fertilization with nitrogen (N) or phosphorus (P). Leachate C:N varied approximately twofold, and C:P varied nearly 20-fold across species and nutrient treatments. Uptake of DOC was greatest for leachates derived from trees that had been fertilized with P, a finding consistent with P-limitation of uptake and/or preferential sorption of P-containing molecules. Optical measures indicated that leachates derived from the three tree species varied in molecular composition, but uptake of DOC did not differ across species, suggesting weak constraints on retention imposed by molecular composition relative to nutrient limitation. Observed coupling between P and C cycles highlights the potential for increased P availability to enhance DOC retention in headwater streams.
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Lignin impedes access to cellulose during biofuel production and pulping but trees can be genetically modified to improve processing efficiency. Modification of lignin may have nontarget effects on mechanical and chemical resistance and subsequent arthropod community responses with respect to pest susceptibility and arthropod biodiversity. We quantified foliar mechanical and chemical resistance traits in lignin-modified and wild-type (WT) poplar (Populus alba × Populus tremula) grown in a plantation and censused arthropods present on these trees to determine total abundance, as well as species richness, diversity and community composition. Our results indicate that mechanical resistance was not affected by lignin modification and only one genetic construct resulted in a (modest) change in chemical resistance. Arthropod abundance and community composition were consistent across modified and WT trees, but transgenics produced using one construct exhibited higher species richness and diversity relative to the WT. Our findings indicate that modification of lignin in poplar does not negatively affect herbivore resistance traits or arthropod community response, and may even result in a source of increased genetic diversity in trees and arthropod communities.
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Phenolics are the most abundant plant metabolites and are believed to decompose slowly in soils compared to other soil organic matter (SOM). Thus, they have often been considered as a slow carbon (C) pool in soil dynamics models. Here, however, we review changes in our concept about the turnover rate of phenolics and quantification of different types of phenolics in soils. Also, we synthesize current research on the degradation of phenolics and their regulatory effects on decomposition. Environmental changes, such as elevated CO 2 , warming, nitrogen (N) deposition, and drought, could influence the production and form of phenolics, leading to a change in SOM dynamics, and thus we also review the fate of phenolics under environmental disturbances. Finally, we propose the use of phenolics as a tool to control rates of SOM decomposition to stabilize organic carbon in ecosystems. Further studies to clarify the role of phenolics in SOM dynamics should include improving quantification methods, elucidating the relationship between phenolics and soil microorganisms, and determining the interactive effects of combinations of environmental changes on the phenolics production and degradation and subsequent impact on SOM processing.
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Analyses of nitrogen and phosphorus in the senesced leaves of 89 species of deciduous and evergreen woody perennials were used (1) to discover the limits of ultimate potential resorption (maximal withdrawal of nutrients from senescing leaves), (2) to determine a means by which resorption can be categorized as complete or incomplete, (3) to develop the concept of resorption proficiency (measured as the levels to which nutrients have been reduced in senesced leaves), (4) to compare resorption in evergreen vs. deciduous species, (5) to assess the impact of phylogeny on resorption, (6) to compare resorption in actinorhizal vs. non-nitrogen-fixing species, and (7) to consider the efficacy of using multiple measures of resorption to answer questions regarding the function and evolution of this process, rather than relying solely on analyses of resorption efficiency (percentage reduction of nutrients between green and senesced leaves). Concentrations of 0.3% nitrogen and 0.01% phosphorus in senesced leaves represent ultimate potential resorption of these nutrients in woody perennials. Resorption proficiency and potential resorption were quantitatively defined in two models that describe both resorption that is maximal and biochemically complete, and that which is not. Resorption is highly proficient in plants that have reduced nitrogen and phosphorus in their senescing leaves to concentrations below 0.7% and 0.05%, respectively. An important feature of knowing the levels to which nutrients can be reduced in senescing leaves is that these values offer an objective gauge by which to measure the success of resorption as a nutrient conservation mechanism. Evergreens were significantly more proficient at resorbing phosphorus than were deciduous species (0.045% vs. 0.067% P in senesced leaves, respectively) and plants capable of symbiotic nitrogen fixation were significantly less proficient at resorbing nitrogen than were nonfixers (1.6% vs. 0.9% N in senesced leaves, respectively). Resorption proficiency appeared to parallel some phylogenic trends, yet the influence of phylogeny was not so significant as to overwhelm the effects of recent selection. The ability of plants to reduce nitrogen in senescing leaves was significantly correlated with their ability to reduce phosphorus. Measurement and analysis of resorption proficiency, when coupled with concurrent consideration of potential resorption and resorption efficiency, should facilitate and expedite the ongoing attempt to resolve complex questions regarding the environmental constraints that influence resorption, and the selection pressures that have directed the evolution of this process.
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Our aim in this study was to determine how well phenotypic variation iri foliar concentrations of carbon-based secondary compounds (CBSCs) in woody plants can be predicted on the basis of two resource-based hypotheses, i.e. the carbon-nutrient balance (CNB) and growth-differentiation balance (GDB) hypotheses. We conducted a meta-analysis of literature data with respect to responses of CBSCs, carbohydrates and nitrogen to six types of environmental manipulations (fertilization with nitrogen or phosphorus, shading, CO2 enrichment, drought stress, ozone exposure): Plant responses to nitrogen fertilization, shading and CO2 enrichment in terms of pooled CBSCs and carbohydrates were consistent with predictions made with the two hypotheses. However, among biosynthetically distinct groups of CBSCs only concentrations of phenylpropanoid-derived compounds changed as predicted; hydrolyzable tannins and terpenoids, in particular, were less responsive. Phosphorus fertilization did not affect concentrations of CBSC or primary metabolites. Plant responses to drought and ozone exposure presumably were driven by plant demands for particular types of compounds (osmolites in the case of drought and antioxidants in the case of ozone exposure) rather than by changes in resource availability. Based on the relative importance of the treatment effects, we propose a hierarchical model of carbon allocation to CBSCs. The model implies that CBSC production is determined by both resource availability and specific demand-side responses. However, these two mechanisms work at different hierarchical levels. The domain of the CNB and GDB hypotheses is at the high hierarchical levels, predicting the total amount of carbon that can be allocated to CBSCs. Predicting altered concentrations of individual CBSCs, i.e. low hierarchy levels, probably demands biosynthetically detailed models which also take into account the history of plant interactions with biotic and abiotic factors.
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From literature sources we compiled the data on carbon-based-secondary compounds CBSC (phenolics and terpenoids) and biomass of 17 plant species grown at different CO2 concentrations under low and high nutrient availabilities. With a low nutrient availability a possible inverse correlation was found between the biomass and CBSC changes. On the contrary, under a high nutrient availability, both the CBSC and biomass increased with elevated CO2. The wide variation in the CBSC production among species and compounds (larger responses in phenolics than in terpenoids) indicates that the allocation to CBSC may not completely be governed by changes in CO2 and nutrient availabilities per se. Yet the comparison shows that elevated CO2 generally loads the carbon into CBSC [their leaf concentration increased an overall average of 14 % at 700 umol(CO2) mol-r] which may improve our understanding of the carbon storage and cycling in ecosystems under the “global change” of climate.
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Increasing concentrations of atmospheric COâ will interact with other environmental factors to influence the physiology and ecology of trees. This research evaluated how plant phytochemical responses to enriched atmospheric COâ are affected by the availability of soil nitrate (NOâ⁻) and how these chemical changes alter performance of a tree-feeding folivore. Seedlings of three deciduous tree species were grown in ambient or elevated COâ in combination with low or high soil NOâ⁻ availability. Lymantria dispar larvae were reared on foliage (aspen and maple). Concentrations of nitrogen and soluble protein decreased, whereas concentrations of starch, condensed tannins, and ellagitannins increased, in response to elevated COâ and/or low NOâ⁻. Responses of simple carbohydrates and phenolic glycosides were variable absolute (net) changes in foliar C:N ratios were greatest for aspen and least for oak, whereas relative changes were greatest for maple and least for aspen. Elevated COâ treatments had little effect on gypsy moth development time, growth rate, or larval mass. Larvae reared on aspen foliage grown under elevated COâ exhibited increased consumption but decreased conversion efficiencies. Gypsy moth responses to NOâ⁻ were strongly host specific. The magnitude of insect response elicited by resource-mediated shifts in host chemistry will depend on how levels of compounds with specific importance to insect fitness are affected. Relatively few true interactions occured between carbon and nitrogen availability and insect performance. Tree species frequently interacted with COâ and/or NOâ⁻ availability to affect both parameters. The effects of elevated atmospheric COâ on terrestrial plant communities will depend on species composition and soil nutrient availability. 54 refs., 9 figs., 4 tabs.
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We suggest that selective foraging alters feedbacks between plants and decomposers and between plants and herbivores. Plant tissue chemistry is an important link between herbivores and decomposers. Plants that produce easily decomposable litter are also heavily browsed, because the same chemical properties that determine litter decay also determine digestibility. This trait links theories of food webs and nutrient cycles by posing a role of herbivores as functional switches determining both plant community composition and the array of litters returned to the soil. This role appears to be particularly strong in boreal forests, where nutrient availability is low and limits productivity and determines successional pathways, where effects of herbivores are strong and long lasting, and where the same plant traits that determine herbivore preference and response to browsing also determine interactions with soil nutrient availability. Such feedbacks cause the effects of herbivores on ecosystems to persist even after the herbivores are no longer present.
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The prediction that litter quality, and hence litter decomposition rates, would be reduced when plants are grown in a CO2-enriched atmosphere has been based on the observation that foliar N concentrations usually are lower in elevated [CO2]. The implicit assumption is that the N concentration in leaf litter reflects the N concentration in green leaves. Here we evaluate that assumption by exploring whether the process of seasonal nutrient resorption is different in CO2-enriched plants. Nitrogen resorption was studied in two species of maple trees (Acer rubrum L. and A. saccharum Marsh.), which were planted in unfertilized soil and grown in open-top chambers with ambient or elevated [CO2] in combination with ambient or elevated temperature. In the second growing season, prior to autumn senescence, individual leaves were collected and analyzed for N and dry matter content. Other leaves at the same and an adjacent node were collected for analysis as they senesced and abscised. This data set was augmented with litter samples from the first growing season and with green leaves and leaf litter collected from white oak (Quercus alba L.) saplings grown in ambient and elevated [CO2] in open-top chambers. In chambers maintained at ambient temperature, CO2 enrichment reduced green leaf N concentrations by 25% in A. rubrum and 19% in A. saccharum. CO2 enrichment did not significantly reduce resorption efficiency so the N concentration also was reduced in litter. There were, however, few effects of [CO2] on N dynamics in these leaves; differences in N concentration usually were the result of increased dry matter content of leaves. The effects of elevated [CO2] on litter N are inherently more difficult to detect than differences in green leaves because factors that affect senescence and resorption increase variability. This is especially so when other environmental factors cause a disruption in the normal progress of resorption, such as in the first year when warming delayed senescence until leaves were killed by an early frost. The results of this experiment support the approach used in ecosystem models in which resorption efficiency is constant in ambient and elevated [CO2], but the results also indicate that other factors can alter resorption efficiency.
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The influence of chemical and physical quality of litter on its rate of decomposition was examined by measuring mass loss from 19 diverse litter types, including leaves, needles, forbs, wood, and roots in the Kananaskis Valley of Alberta, Canada. Litter samples drawn from three adjacent forests of lodgepole pine, white spruce, and Engelmann spruce – subalpine fir, and from a small clearcut area, were allowed to decompose for 3 years at their sites of origin. The best predictors of mass loss were the initial concentrations of lignin and labile material in the litter. Adding N or P contents as a second term, rather than as a lignin to nutrient ratio, significantly improved mass loss predictions. There were abrupt limits for the influence of lignin (above 28%) and N (below C:N of 30:1); similar limits were observed for all predictors except labile content. None of these chemical parameters, nor a physical measure, particle diameter, were useful in predicting rates of decomposition of high-lignin, woody substrates. The relative importance of the various litter quality parameters in determining rates of mass loss in the clear-cut area was very similar to that in the forests, despite considerably more rapid decomposition in the clearcut. Key words: decomposition, lignin, litter, nutrients, Rocky Mountains, wood.
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Leaf litter from 8 species of tree, shrub or herb, ranging in lignin content from 3.4-20.5%, was allowed to decompose in microcosms for up to 4 mo (equivalent to 1.5-2 yr decay in the field). Nitrogen content and the C:N ratio were the best predictors of mass loss rate, and were substantially better than the ligin:N ratio. However, when regressions were tested using pine needles (lignin content 26.2%), the C:N ratio and N content badly underestimated mass remaining, while lignin content and the lignin:N ratio overestimated it by <2%. Regressions of initial lignin content or lignin:N ratio on mass remaining improved from 2 to 4 mo decomposition, while those of N content grew worse, illustrating succession of N to lignin control of decomposition rate. -from Authors
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This study was conducted to examine the effects of CO2-mediated changes in tree chemistry on the performance of the gypsy moth ((Lymantria dispar L.) and the parasitold Cotesia melanoscela (Ratz.). We used carbon-nutrient balance theory to develop hypotheses regarding changes in tree chemistry and the performance of both insects under elevated CO2. As predicted, levels of foliar nitrogen declined and concentrations of carbon-based compounds (e.g. starch and phenolics) increased under elevated CO2. Gypsy moth performance (e.g. growth, development) was altered by CO2-mediated changes in foliar chemistry, but the magnitude was small and varied across tree species. Larvae feeding on high CO2 aspen exhibited the largest reduction in performance, relative to larvae feeding on birch, oak, or maple. Parasitism by C. melanoscela significantly prolonged gypsy moth development and reduced growth rates. Overall, the effect of parasitism on gypsy moth performance did not differ between CO2 treatments. Altered gypsy moth performance on high CO2 foliage in turn affected parasitoid performance, but the response was variable: parasitoid mortality increased and adult female size declined slightly under high CO2, while development time and adult male size were unaffected. Our results suggest that CO2-induced changes in plant chemistry were buffered to the extent that effects on third trophic level interactions were weak to non-existent for the system examined in this study.
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Enriched atmospheric CO2 alters the quantity and quality of plant production, but how such effects vary among plant genotypes is poorly known. We evaluated the independent and interactive effects of CO2 and nutrient availability on growth, allocation and phytochemistry of six aspen (Populus tremuloides Michx.) genotypes. One-year-old trees, propagated from root cuttings, were grown in CO2-controlled glasshouses for 64 days, then harvested. Foliage was analyzed for levels of water, nitrogen, starch, phenolic glycosides and condensed tannins. Of seven plant growth/allocation variables measured, four (biomass production, stem growth, relative growth rate and root:shoot ratio) exhibited marginally to highly significant CO2 2 genotype interactions. CO2 enrichment stimulated growth of some genotypes more than others, and this interaction was itself influenced by soil nutrient availability. In addition, enriched CO2 increased the magnitude of the among-genotype variance for four of the growth/allocation variables. Of six foliar chemical constituents analyzed, CO2-mediated responses of two (the phenolic glycoside tremulacin and condensed tannins) varied among genotypes. Moreover, enriched CO2 increased the magnitude of among-genotype variance for four of the chemical variables. Given the importance of these growth and chemical characteristics to the biological fitness of aspen, this research suggests that projected atmospheric CO2 increases are likely to alter the genetic structures and evolutionary trajectories of aspen populations.
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Plant species differ broadly in their responses to an elevated CO2 atmosphere, particularly in the extent of nitrogen dilution of leaf tissue. Insect herbivores are often limited by the availability of nutrients, such as nitrogen, in their host plant tissue and may therefore respond differentially on different plant species grown in CO2-enriched environments. We reared gyspy moth larvae (Lymantria dispar) in situ on seedlings of yellow birch (Betula allegheniensis) and gray birch (B. populifolia) grown in an ambient (350 ppm) or elevated (700 ppm) CO2 atmosphere to test whether larval responses in the elevated CO2 atmosphere were species-dependent. We report that female gypsy moths (Lymantria dispar) reared on gray birch (Betula populifolia) achieved similar pupal masses on plants grown at an ambient or an elevated CO2 concentration. However, on yellow birch (B. allegheniensis), female pupal mass was 38% smaller on plants in the elevated-CO2 atmosphere. Larval mortality was significantly higher on yellow birch than gray birch, but did not differ between the CO2 treatments. Relative growth rate declined more in the elevated CO2 atmosphere for larvae on yellow birch than for those on gray birch. In preference tests, larvae preferred ambient over elevated CO2-grown leaves of yellow birch, but showed no preference between gray birch leaves from the two CO2 atmospheres. This differential response of gypsy moths to their host species corresponded to a greater decline in leaf nutritional quality in the elevated CO2 atmosphere in yellow birch than in gray birch. Leaf nitrogen content of yellow birch dropped from 2.68% to 1.99% while that of gray birch leaves only declined from 3.23% to 2.63%. Meanwhile, leaf condensed tannin concentration increased from 8.92% to 11.45% in yellow birch leaves while gray birch leaves only increased from 10.72% to 12.34%. Thus the declines in larval performance in a future atmosphere may be substantial and host-species-specific.
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Increases in tree biomass may be an important sink for CO2 as the atmospheric concentration continues to increase. Tree growth in temperate forests is often limited by the availability of soil nutrients. To assess whether soil nutrient limitation will constrain forest productivity under high atmospheric CO2, we studied the changes in forest litter production and nutrient cycling in a maturing southern U.S. loblolly pine-hardwood forest during two years of free-air CO2 enrichment. The objective of this paper is to present data on the chemistry of green leaves and leaf litter, nutrient-retranslocation efficiency, above-ground litter production, whole-system nutrient-use efficiency, decomposition, and N availability in response to forest growth under elevated CO2. The chemical composition of green leaves and leaf litter was largely unaffected by elevated CO2. Green-leaf nitrogen (N) and phosphorus (P) concentrations were not significantly lower under elevated CO2. N and P retranslocation from green leaves did not increase under elevated CO2; therefore, leaf litter N and P concentrations were not significantly lower under elevated CO2. The concentrations of carbon, lignin, and total nonstructural carbohydrates in litter were not significantly different under elevated CO2. Total aboveground litterfall increased significantly with CO2 fumigation. The increase in litterfall was due to significant increases in loblolly pine leaf litter and bark production. The mass of leaves from deciduous species did not increase with CO2 fumigation. Whole-system nutrient-use efficiency (aboveground litterfall/nutrient content of litterfall) did not increase as a consequence of forest growth under elevated CO2, but N and P fluxes from vegetation to the forest floor increased significantly. During the second year of CO2 fumigation, the flux of N and P to the forest floor in litterfall increased by 20% and 34%, respectively. The rate of mass loss during one year of decomposition was unaffected by "litter type" (whether the litter was produced under ambient or elevated CO2), nor by the "site" of decomposition (whether the litter was decomposed in the ambient or elevated CO2 plots). N was immobilized in litter during decomposition, whereas P was mineralized. There was no consistent effect of litter type or site on nutrient dynamics in decomposing litter. There was no significant effect of elevated CO2 on the pool size of inorganic N (NH4 + and NO3 -) in the top 7.5 cm of mineral soil. The rate of net N mineralization and nitrification in mineral soil was not significantly different between treatment and control plots. Identifying the source of the nutrients lost in litterfall is critical to the long-term potential growth stimulation of forests under elevated CO2. If the nutrients lost from biomass come from storage (e.g., the movement of nutrients from wood to leaves), then the increase in litter production should decrease over time as slowly replenished nutrient reserves are exhausted. If the nutrients lost in plant litter are replaced by uptake from soils, then it is possible (1) that trees acquire soil nutrients at a rate commensurate with growth stimulated by elevated CO2; and (2) that forest productivity will be stimulated by elevated CO2 in the long term.
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This study examined the effects of CO2 and light availability on sapling growth and foliar chemistry, and consequences for insect performance. Quaking aspen (Populus tremuloides Michx.), paper birch (Betula papyrifera Marsh.), and sugar maple (Acer saccharum Marsh.) were grown in controlled environment greenhouses under ambient or elevated CO2 (38.7 and 69.6 Pa), and low or high light availability (375 and 855 μmol m−2 s−1). Because CO2 and light are both required for carbon assimilation, the levels of these two resources are expected to have strong interactive effects on tree growth and secondary metabolism. Results from this study support that prediction, indicating that the relative effect of rising atmospheric CO2 concentrations on the growth and secondary metabolism of deciduous trees may be dependent on light environment. Trees in ambient CO2-low light environments had substantial levels of phytochemicals despite low growth rates; the concept of basal secondary metabolism is proposed to explain allocation to secondary metabolites under growth-limiting conditions. Differences between CO2 and light effects on the responses of growth and secondary metabolite levels suggest that relative allocation is not dependent solely on the amount of carbon assimilated. The relative growth rates and indices of feeding efficiency for gypsy moth (Lymantria dispar L.) larvae fed foliage from the experimental treatments showed no significant interactive effects of light and CO2, although some main effects and many host species interactions were significant. Gypsy moth performance was negatively correlated with CO2- and light-induced increases in the phenolic glycoside content of aspen foliage. Insects were not strongly affected, however, by treatment differences in the nutritional and secondary chemical components of birch and maple.
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Current models of climate change predict a reduction of area covered by northern coniferous forests and tundra, and an increase in grasslands. These scenarios also indicate a northerly shift in agricultural regions, bringing virgin soils under cultivation. The direct effects of man on tundra, boreal forest, and temperate grassland ecosystems are likely to result in less carbon mobilization from soils and vegetation than from tropical forests. However, as a consequence of climate change, carbon mineralization rates from arctic and sub-arctic soils could be very rapid under warmer and drier conditions because of low stabilization of soil organic matter (SOM) and enhanced microbial responses to small changes in soil moisture and temperature. Predicting the response of these systems to climate change is complicated where the edaphic environment regulating SOM dynamics is not a direct function of macroclimatic conditions. The primary recommendation for future research is for integrated studies on plant and soil processes. -from Author
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1. Interactions between trees and tree-feeding insects are likely to shift under conditions of enriched atmospheric CO<sub>2</sub> owing to changes in foliar chemical composition. This study addressed the effects of CO<sub>2</sub>-mediated changes in leaf chemistry on performance of three silkmoth (Saturniidae) species: cecropia (Hyalophora cecropia), luna (Actias luna) and polyphemus (Antheraea polyphemus polyphemus). 2. Growth under elevated CO<sub>2</sub> atmospheres decreased nitrogen concentrations (23%) but tripled starch and doubled condensed tannin concentrations, resulting in a marked increase in foliar carbon:nitrogen ratio. 3. Survival of first stadium larvae was marginally reduced when reared on high CO<sub>2</sub> leaves. 4. Development rates were prolonged, growth rates tended to decline, consumption increased and food processing efficiencies decreased for fourth stadium larvae reared on high CO<sub>2</sub> leaves. The magnitude of responses varied among species. 5. Overall performance of these saturniid species, at least when feeding on birch, is predicted to decline under atmospheric CO<sub>2</sub> conditions anticipated for the next century.
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The evolutionary response of plants to herbivory is constrained by the availability of resources in the environment. Woody plants adapted to low-resource environments have intrinsically slow growth rates that limit their capacity to grow rapidly beyond the reach of most browsing mammals. Their low capacity to acquire resources limits their potential for compensatory growth which would otherwise enable them to replace tissue destroyed by browsing. Plants adapted to low-resource environments have responded to browsing by evolving strong constitutive defenses with relatively low ontogenetic plasticity. Because nutrients are often more limiting than light in boreal forests, slowly growing boreal forest trees utilize carbon-based rather than nitrogen-based defenses. More rapidly growing shade-intolerant trees that are adapted to growth in high-resource environments are selected for competitive ability and can grow rapidly beyond the range of most browsing mammals. Moreover, these plants have the carbon and nutrient reserves necessary to replace tissue lost to browsing through compensatory growth. However, because browsing of juvenile plants reduces vertical growth and thus competitive ability, these plants are selected for resistance to browsing during the juvenile growth phase. Consequently, early successional boreal forest trees have responded to browsing by evolving strong defenses during juvenility only. Because severe pruning causes woody plants to revert to a juvenile form, resistance of woody plants to hares increases after severe hare browsing as occurs during hare population outbreaks. This increase in browsing resistance may play a significant role in boreal forest plant-hare interactions. Unlike woody plants, graminoids retain large reserves of carbon and nutrients below ground in both low-resource and high-resource environments and can respond to severe grazing through compensatory growth. These fundamental differences between the response of woody plants and graminoids to vertebrate herbivory suggest that the dynamics of browsing systems and grazing systems are qualitatively different. /// Эволюционная реакция растений на выедание фитофагами органичивается дос-тупностью ресурсов в природной среде. Древесые растения, адаптированные к местообитаниям с низкнм количеством ресурсов, имеют эндогенные низкие темпы роста, что лимитирует их способность к быстрому росту после повреж-дения мпекопитающими, обьедающими побеги. Их низкая способность к утили-зации ресурсов ограничивает их потенцию к компенсаторному росту, возме-щающему ткани, поврежденные при обьедании побегов. Растения, адаптирован-ные к местообитаниям с низким запасом ресурсов, реагируют на повреждения развитием сильных конструктивных защитных механизмов с относительно низ-кой онтогенетической пластичностью. Так как элементы питания часто более ограничены, чем освещение в бореальных лесах, медленно растущие деревья бореальных лесов используют защитные механизмы, основанные чаще на угле-родных, нежели азотных соединениях. У быстрее растущих деревьев, чувстви-тельных к затенеию, которые адаптированы к росту в более обеспеченных ре-сурсами местообитаниях, отбор был направлен на конкурентоспособность, они могут быстро расти после повреждений большинства млекопитакщих, обьедаю-щих побеги. Голее того, эти растения имеют резервы углерда и азота, не-обходимые для замещения обьеденных тканей в процессе компесаторного роста. Однако, так как обьедание молодых растений снижает их вертикальный рост и таким образом конкурентоспособность у этих растений отбор направлен на устойчивость к обьеданию в течение ювенильной фазы роста. Следовательно, деревья бореальных лесов ранний сукцессионной реагируют на обьеда-ние рзвитием сильных защитных механизмов лишь в незрелом состоянии. Так как сильное обьедание возращает древесные растения к ювенильной форме, у-стойчивость древесных растений к повреждениям зайцев поьшается после сильного обьедания зайцами, как это наблюдалось в периоды вспышек числен-ности их популяций. Это повышение устойчивости к обьеданию может играть существенную роль в отношениях растение-заяц, в бореальных лесах. В отли-чие от древесных растений, злаковые сохраняют резервы углерода и других элементов в подземных частях в местообитаниях с низкой и высокой обеспе-ченностью ресурсами и могут реагировать на сильный выпас компенсаторным ростом. Эти функциональные различия между реакцией древесных растений и злаков на выедание млекопитаюшими показывает, что динамика систем обгла-дывания и пастьбы качественно различны.
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Six years (1978-1983) of ozone monitoring data from sites located within six forested areas were examined. Areas that experienced the lowest to the highest ozone exposures were located in (1) northern New England/New York and upper Great Lakes, (2) New York/Pennsylvania/Maryland, (3) southeastern/southern, and (4) New Jersey pinelands. In general, higher ozone concentrations were observed in 1978, 1980 and 1983 as compared to the other three years examined. Ozone concentrations varied considerably within the areas. Recommendations for additional ozone monitoring sites are made. A concentrated effort should be made to examine ozone monitoring data from subsequent years (1984, 1985, and 1986) to explore whether the 6-year period 1978 through 1983 is representative of the annual variability of ozone concentrations over eastern forested areas. To better understand the relationship between ozone exposure and possible forest effects, we recommend that the temporal distributions of elevated ozone concentrations over a growing season be examined. The occurrence of elevated ozone levels during specific growth periods during a season may be an important aspect that biologists may wish to explore.
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The carbon/nutrient balance hypothesis suggests that leaf carbon to nitrogen ratios influence the synthesis of secondary compounds such as condensed tannins. We studied the effects of rising atmospheric carbon dioxide on carbon to nitrogen ratios and tannin production. Six genotypes of Populus tremuloides were grown under elevated and ambient CO 2 partial pressure and high- and low-fertility soil in field open-top chambers in northern lower Michigan, USA. During the second year of exposure, leaves were harvested three times (June, August, and September) and analyzed for condensed tannin concentration. The carbon/nutrient balance hypothesis was supported overall, with significantly greater leaf tannin concentration at high CO2 and low soil fertility compared to ambient CO2 and high soil fertility. However, some genotypes increased tannin concentration at elevated compared to ambient CO2, while others showed no CO2 response. Performance of lepidopteran leaf miner (Phyllonorycter tremuloidiella) larvae feeding on these plants varied across genotypes, CO2, and fertility treatments. These results suggest that with rising atmospheric CO 2, plant secondary compound production may vary within species. This could have consequences for plant‐herbivore and plant‐microbe interactions and for the evolutionary response of this species to global climate change.
Article
Direct and interactive effects of CO 2 and light on tree phytochemistry and insect fitness parameters were examined through experimental manipulations of plant growth conditions and performance of insect bioassays. Three species of deciduous trees (quaking aspen, Populus tremuloides ; paper birch, Betula papyrifera ; sugar maple, Acer saccharum ) were grown under ambient (387±8 μL/L) and elevated (696±2 μL/L) levels of atmospheric CO 2 , with low and high light availability (375 and 855 μmol×m ⁻² ×s ⁻¹ at solar noon). Effects on the population and individual performance of a generalist phytophagous insect, the white‐marked tussock moth ( Orgyia leucostigma ) were evaluated. Caterpillars were reared on experimental trees for the duration of the larval stage, and complementary short‐term (fourth instar) feeding trials were conducted with insects fed detached leaves. Phytochemical analyses demonstrated strong effects of both CO 2 and light on all foliar nutritional variables (water, starch and nitrogen). For all species, enriched CO 2 decreased water content and increased starch content, especially under high light conditions. High CO 2 availability reduced levels of foliar nitrogen, but effects were species specific and most pronounced for high light aspen and birch. Analyses of secondary plant compounds revealed that levels of phenolic glycosides (salicortin and tremulacin) in aspen and condensed tannins in birch and maple were positively influenced by levels of both CO 2 and light. In contrast, levels of condensed tannins in aspen were primarily affected by light, whereas levels of ellagitannins and gallotannins in maple responded to light and CO 2 , respectively. The long‐term bioassays showed strong treatment effects on survival, development time, and pupal mass. In general, CO 2 effects were pronounced in high light and decreased along the gradient aspen birch maple. For larvae reared on high light aspen, enriched CO 2 resulted in 62% fewer survivors, with increased development time, and reduced pupal mass. For maple‐fed insects, elevated CO 2 levels had negative effects on survival and pupal mass in low light. For birch, the only negative CO 2 effects were observed in high light, where female larvae showed prolonged development. Fourth instar feeding trials demonstrated that low food conversion efficiency reduced insect performance. Elevated levels of CO 2 significantly reduced total consumption, especially by insects on high light aspen and low light maple. This research demonstrates that effects of CO 2 on phytochemistry and insect performance can be strongly light‐dependent, and that plant responses to these two environmental variables differ among species. Overall, increased CO 2 availability appeared to increase the defensive capacity of early‐successional species primarily under high light conditions, and of late‐successional species under low light conditions. Due to the interactive effects of tree species, light, CO 2 , and herbivory, community composition of forests may change in the future.
Article
In field experiments three clones of two-year-old birch (Betula pendula Roth.) were exposed for one growing season to an ozone concentration 1–2 times higher than ambient. At the end of August, leaf and stem material was analyzed for a wide range of primary and secondary metabolites. Although most of these metabolites were not significantly affected by ozone exposure, ozone-treated leaves contained larger concentrations of total sugars and reduced amounts of the phenolic glucoside, dehydrosalidroside. In the stems, greater amounts of catechin pentoside, hyperoside and papyriferic acid were found. The results indicated considerable inter- and intra-clonal variation in the production of phytochemicals in both leaves and stems.
Article
A simple and sensitive method for the colorimetric determination of reducing sugars in plant materials is proposed. The procedure is based on reduction under alkaline conditions of potassium ferricyanide by the reducing groups of the carbohydrates, followed by colour development as the o-phenanthroline complex. All sugars tested produced an equal colour yield. The method proved to be reproducible and precise with a high degree of sensitivity which makes it possible to use with small sample quantities. The method is quick, easy and cheap, hence permiting its use in routine analysis. © 1998 John Wiley & Sons, Ltd.
Article
We review experimental studies to evaluate how the nitrogen cycle influences the response of forest net primary production (NPP) to elevated CO2. The studies in our survey report that at the tissue level, elevated CO2 reduces leaf nitrogen concentration an average 21%, but that it has a smaller effect on nitrogen concentrations in stems and fine roots. In contrast, higher soil nitrogen availability generally increases leaf nitrogen concentration. Among studies that manipulate both soil nitrogen availability and atmospheric C02, photosynthetic response depends on a linear relationship with the response of leaf nitrogen concentration and the amount of change in atmospheric CO2 concentration. Although elevated CO2 often results in reduced tissue respiration rate per unit biomass, the link to changes in tissue nitrogen concentration is not well studied. 1The US government has the right to retain a nonexclusive, royalty-free license in and to any copyright covering this paper.
Article
This text is designed for an introductory graduate level course in statistics and experimental design. While the text is oriented toward the behavioral sciences, it may be applicable to other settings as well—an undergraduate class in applied statistics or the medical sciences. Chapters 1 and 2 cover elementary probability and statistical inference, while Chapters 3 and 4 deal with the general linear model and the general linear hypothesis. Chapter 5 is an overview of experimental design. Chapters 6 through 8 contain more than 40 example applications of the univariate normal, multivariate normal, and multinomial analysis of variance models. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Article
In the first part of this review, I discuss how we can predict the direct short-term effect of enhanced CO2 on photosynthetic rate in C3 terrestrial plants. To do this, I consider: (1) to what extent enhanced CO2 will stimulate or relieve demand on partial processes like carboxylation, light harvesting and electron transport, the Calvin cycle, and end-product synthesis; and (2) the extent to which these various processes actually control the rate of photosynthesis. I conclude that control is usually shared between Rubisco (which responds sensitively to CO2) and other components (which respond less sensitively), and that photosynthesis will be stimulated by 25–75% when the CO2 concentration is doubled from 35 to 70 Pa. This is in good agreement with the published responses. In the next part of the review, I discuss the evidence that most plants undergo a gradual inhibition of photosynthesis during acclimation to enhanced CO2. I argue that this is related to an inadequate demand for carbohydrate in the remainder of the plant. Differences in the long-term response to CO2 may be explained by differences in the sink-source status of plants, depending upon the species, the developmental stage, and the developmental conditions. In the third part of the review, I consider the biochemical mechanisms which are involved in ‘sink’ regulation of photosynthesis. Accumulating carbohydrate could lead to a direct inhibition of photosynthesis, involving mechanical damage by large starch grains or Pi-limitation due to inhibition of sucrose synthesis. I argue that Pi is important in the short-term regulation of partitioning to sucrose and starch, but that its contribution to ‘sink’ regulation has not yet been conclusively demonstrated. Indirect or ‘adaptive’ regulation of photosynthesis is probably more important, involving decreases in amounts of key photosynthetic enzymes, including Rubisco. This decreases the rate of photosynthesis, and potentially would allow resources (e.g. amino acids) to be remobilized from the leaves and reinvested in sink growth to readjust the sink-source balance. In the final part of the review, I argue that similar changes of Rubisco and, possibly, other proteins are probably also involved during acclimation to high CO2.
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We summarize the impacts of elevated CO2 on the N concentration of plant tissues and present data to support the hypothesis that reductions in the quality of plant tissue commonly occur when plants are grown under elevated CO2. Synthesis of existing data showed an average 14% reduction of N concentrations in plant tissue generated under elevated CO2 regimes. However, elevated CO2 appeared to have different effects on the N concentrations of different plant types, as the reported reductions in N have been larger in C3 plants than in C4 plants and N2-fixers. Under elevated CO2 plants changed their allocation of N between above- and below-ground components: root N concentrations were reduced by an average of 9% compared to a 14% average reduction for above-ground tissues. Although the concentration of CO2 treatments represented a significant source of variance for plant N concentration, no consistent trends were observed between them.
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Current estimates of forest yield losses attributable to ozone pollution amount to c . 10% over Europe as a whole. This figure is derived from a synthesis of all European studies using trees for which AOT40 exposure values are available. However, the choice of 40 nl l ⁻¹ as the threshold concentration for demonstrable effects has led to debate, and this value might not be low enough to predict ozone effects in Scandinavia, where concentrations are lower than in southern Europe, and chronic injury resulting from cumulative exposure is observed. This ‘level I’ approach provides a useful means for mapping physiologically effective concentrations but has significant shortcomings in that it is unable to take environmental conditions into account. In order to produce a mechanism capable of predicting yield losses resulting from ozone pollution at specific sites and for individual species, the effective ozone dose, a product of conductance and concentration, must be calculated. Process‐based models relating the environment (temperature, humidity/saturation deficit, incident light and soil moisture content) to conductance are available for a number of species (oak, Scots pine, Norway spruce, Sitka spruce, beech and poplar). Effective ozone doses could therefore be calculated, and the relationships between effective dose and yield loss could be determined by revisiting existing data for which only concentrations or AOT40 values are available at present. Yield loss must be the growth parameter with which ozone damage is expressed, since visible injury does not necessarily represent severe injury to the plant, whilst visibly unaffected plants may be significantly compromised in terms of biomass accumulation. For conifers, premature needle drop, although indicative of ozone pollution, might not represent a significant reduction in growth since older needles are, functionally, relatively unimportant. When revisiting old experiments the question should also be asked ‘what is an appropriate control treatment’. It is unrealistic to expect ambient O 3 concentrations on a global scale to return to pre‐industrialization levels, and therefore the use of a charcoal‐filtered treatment is probably unrealistic. In addition, charcoal filters will remove some of the NO x , and therefore on nutrient deficient soils (typical of many forest soils) the ‘control’ treatment might represent a reduced supply of nitrogen, making it difficult to disentangle the ozone effect per se . The stage has therefore been reached where ‘level II’ ozone exposure mapping (i.e. based on effective dose at the physiological level) is possible for a number of species. As financial considerations become increasingly important, the scientific community must aim to provide better estimates of O 3 ‐induced yield losses so that long‐term environmental audits can be performed.
Article
Quantitative integration of the literature on the effect of elevated CO2 on woody plants is important to aid our understanding of forest health in coming decades and to better predict terrestrial feedbacks on the global carbon cycle. We used meta-analytic methods to summarize and interpret more than 500 reports of effects of elevated CO2 on woody plant biomass accumulation and partitioning, gas exchange, and leaf nitrogen and starch content. The CO2 effect size metric we used was the log-transformed ratio of elevated compared to ambient response means weighted by the inverse of the variance of the log ratio. Variation in effect size among studies was partitioned according to the presence of interacting stress factors, length of CO2 exposure, functional group status, pot size, and type of CO2 exposure facility. Both total biomass (W T) and net CO2 assimilation (A) increased significantly at about twice ambient CO2, regardless of growth conditions. Low soil nutrient availability reduced the CO2 stimulation of W T by half, from +31% under optimal conditions to +16%, while low light increased the response to +52%. We found no significant shifts in biomass allocation under high CO2. Interacting stress factors had no effect on the magnitude of responses of A to CO2, although plants grown in growth chambers had significantly lower responses (+19%) than those grown in greenhouses or in open-top chambers (+54%). We found no consistent evidence for photosynthetic acclimation to CO2 enrichment except in trees grown in pots <0.5 l (−36%) and no significant CO2 effect on stomatal conductance. Both leaf dark respiration and leaf nitrogen were significantly reduced under elevated CO2 (−18% and −16% respectively, data expressed on a leaf mass basis), while leaf starch content increased significantly except in low nutrient grown gymnosperms. Our results provide robust, statistically defensible estimates of elevated CO2 effect sizes against which new results may be compared or for use in forest and climate model parameterization.
Article
Scots pine (Pinus sylvestris L.) trees, aged about 20 years old, growing on a natural pine heath were exposed to two concentrations of CO2 (ambient CO2 and double-ambient CO2) and two O3 regimes (ambient O3 and double-ambient O3) and their combination in open-top chambers during growing seasons 1994, 1995 and 1996. Concentrations of foliar starch and secondary compounds are reported in this paper. Starch concentrations remained unaffected by elevated CO2 and/or O3 concentrations during the first 2 study years. But in the autumn of the last study year, a significantly higher concentration of starch was found in current-year needles of trees exposed to elevated CO2 compared with ambient air. There were large differences in concentrations of starch and secondary compounds between individual trees. Elevated concentrations of CO2 and/or O3 did not have any significant effects on the concentrations of foliar total monoterpenes, total resin acids or total phenolics. Significantly higher concentrations of monoterpenes and resin acids and mostly lower concentrations of starch were found in trees growing without chambers than in those growing in open-top chambers, while there were no differences in concentrations of total phenolics between trees growing without or in chambers. The results suggest that elevated concentrations of CO2 might increase foliar starch concentrations in Scots pine, while secondary metabolites remain unaffected. Realistically elevated O3 concentrations do not have clear effects on carbon allocation to starch and secondary compounds even after 3 exposure years.
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The hydrolysis of proanthocyanidins to anthocyanidins in n-BuOH-HCl (95:5) has been shown to be an autoxidation, the yield of anthocyanidin being critically dependent on trace metal-ion impurities. Reproducible yields of anthocyanidin may be achieved if iron (III) salts are added to the reaction medium, and a standard method of analysis of proanthocyanidins based on use of an n-BuOH-HCl-FeIII mixture is given. The ratio of absorbance maxima of the cyanidin (550 nm) produced to that near 280 nm for the original procyanidin polymer solution was ∼ 3.5.
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We documented temporal patterns in phytochemical composition of Populus tremuloides Michx. during leaf senescence, and the influence of genotype and soil nutrient availability on such patterns. Levels of foliar nitrogen, carbohydrates, phenolic glycosides and condensed tannins were quantified for four aspen genotypes grown in a common garden, with low and high levels of soil nutrients. Levels of all compounds tended to decline over time, although the magnitude of change was influenced by plant genotype and nutrient availability. Genetic variation in concentrations of phytochemicals was much greater for phenolic glycosides and tannins than for nitrogen and carbohydrates, and these phenolic signatures generally persisted through leaf abscission. Our results suggest that genotypic and nutrient effects on patterns of chemical change during senescence will likely influence the performance of late-season herbivores on aspen. Moreover, nutrient and especially genotypic variation in phytochemistry of abscised leaves is likely to affect litter decomposition rates.
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Physiological and ecological constraints play key roles in the evolution of plant growth patterns, especially in relation to defenses against herbivores. Phenotypic and life history theories are unified within the growth-differentiation balance (GDB) framework, forming an integrated system Of theories explaining and predicting patterns of plant defense and competitive interactions in ecological and evolutionary time. Plant activity at the cellular level can be classified as growth (cell division and enlargement) of differentiation (chemical and morphological changes leading to cell maturation and specialization). The GDB hypothesis of plant defense is premised upon a physiological trade-off between growth and differentiation processes. The trade-off between growth and defense exists because secondary metabolism and structural reinforcement are physiologically constrained in dividing and enlarging cells, and because they divert resources from the production of new leaf area. Hence the dilemma of plants:
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It has generally been assumed that increasing atmospheric CO2 concentrations will increase plant carbon-based secondary or structural compounds concentrations. These changes may have far-reaching consequences for herbivory and plant litter decomposition. Recent experimental results provide evidence of increases in concentrations of soluble phenolics and condensed tannins but not in lignin, structural polysaccharides or terpenes. They also show significant effects of these plant chemical changes on herbivores and little or any effects on decomposition. However, there is no consistent evidence of any of these effects at the complex long-term ecosystem level.
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Seedlings of European white birch (Betula pendula Roth) were grown in growth chambers for one growth season under four carbon dioxide regimes (350, 700, 1050 and 1400 ppm) and at three fertilization levels (0, 100 and 500 kg ha−1 monthly). The soluble carbohydrates and secondary phenolics in the leaves and stems were analysed. It was found that fertilizer addition reduced the amounts of glucose and fructose while sucrose remained almost unaffected. The sugar content of leaves increased at 700 ppm and 1050 ppm of CO2 and decreased at the highest CO2 concentration (1400 ppm). The amounts of proanthocyanidins and flavonoids in leaves decreased with fertilization addition and increased with CO2 enrichment. The production of simple phenolic glucosides varied according to the fertilization and CO2 treatments. The triterpenoid content of stems seemed to increase with fertilization and CO2-addition. Our results indicate that the production of phytochemicals in the birch seedlings is very sensitive to both fertilization and CO2 addition, which is in agreement with earlier studies, and thus provide some support for the hypothesis of carbon allocation to plant defence when there is an excess of carbon and nutrient. The considerable variation in the production of secondary components may indicate that the synthesis of these defensive metabolites can be regulated by a plant to certain extent, depending on the ability of the plant to acclimate to changes in the physical environment.
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The effects of single-season tropospheric ozone (O3) exposures on growth, leaf abscission, and biomass of trembling aspen (Populustremuloides Michx.) rooted cuttings and seedlings were studied. Plants were grown in the Upper Peninsula of Michigan in open-top chambers with O3 exposures that ranged from 7 to 92 ppm-h. Depending on the genotype, total seasonal O3 exposure in the range of 50–92 ppm-h had negative impacts on stem, retained leaf, and root biomass accumulation and on diameter growth. Leaf abscission generally increased with increasing O3 exposure and was the principal cause of the decrease in leaf biomass of the O3-treated plants. Considerable genetic variation in O3 responses occurred, as shown by differences in sensitivities among clones and among seedlings. However, the responses to O3 of rooted cuttings and seedlings were similar when seedling means were compared with clonal means for leaf abscission, diameter growth, retained leaf biomass, and root biomass. Comparison of a single square-wave treatment (52 ppm-h) with 70 and 92 ppm-h episodic exposures suggested that the plant response to the square-wave exposure was similar to the response to the highest episodic exposure even though the 92 ppm-h episodic exposure was almost twice the square-wave exposure. Our results are consistent with previous studies that show that P. tremuloides is highly responsive to O3 exposure and this response has a strong genetic component.
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Nutrient movements in the senescing foliage of a Rhode Island aspen were measured. Mean resorption of N, P, and Cu was 43, 51 and 10%, respectively. Aluminium, B, Ca, Fe, Mg, Mn, and Zn increased or remained unchanged in senescing foliage during 1986. Resorption of N and P decreased, respectively, from 56 and 64% in 1986 to 24 and 38% in 1988. Resorption of N and P decreased, respectively, from 56 and 64% in 1986 to 24 and 38% in 1988. Older, larger ramets resorbed less N and Cu. Timing of abscission strongly influenced resorption and may have been related to drought conditions in 1988 and to differential exposure to wind in all years. Resorption of N, P and Cu was lowest in those ramets that lost their leaves earliest and in leaves that senesced earliest on individual ramets. -from Authors
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Abstract We examined the effects of CO2 and defoliation on tree chemistry and performance of the forest tent caterpillar, Malacosoma disstria. Quaking aspen (Populus tremuloides) and sugar maple (Acer saccharum) trees were grown in open-top chambers under ambient or elevated concentrations of CO2. During the second year of growth, half of the trees were exposed to free-feeding forest tent caterpillars, while the remaining trees served as nondefoliated controls. Foliage was collected weekly for phytochemical analysis. Insect performance was evaluated on foliage from each of the treatments. At the sampling date coincident with insect bioassays, levels of foliar nitrogen and starch were lower and higher, respectively, in high CO2 foliage, and this trend persisted throughout the study. CO2-mediated increases in secondary compounds were observed for condensed tannins in aspen and gallotannins in maple. Defoliation reduced levels of water and nitrogen in aspen but had no effect on primary metabolites in maple. Similarly, defoliation induced accumulations of secondary compounds in aspen but not in maple. Larvae fed foliage from the enriched CO2 or defoliated treatments exhibited reduced growth and food processing efficiencies, relative to larvae on ambient CO2 or nondefoliated diets, but the patterns were host species-specific. Overall, CO2 and defoliation appeared to exert independent effects on foliar chemistry and forest tent caterpillar performance.
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The conventional isolation method has been modified in order to minimize protein contamination of tannin purified from high tannin sorghum. The two unique steps of the new procedure are preliminary extraction of the ground grain with ethanol and treatment of the partially purified tannin with phenol to remove traces of noncovalently bound protein. Tannin-associated protein removed by phenol treatment is not a random mixture of all the seed proteins, but consists of several discrete components which have been isolated and partially characterized. These proteins are quite hydrophobic, and one is rich in proline. With only minor changes, the purification method can be used to isolate tannin from seeds of other plants such as legumes.
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The Intergovernmental Panel of Climate Change (IPCC) has concluded that the greenhouse gases carbon dioxide (CO2) and tropospheric ozone (O3) are increasing concomitantly globally. Little is known about the effect of these interacting gases on growth, survival, and productivity of forest ecosystems. In this study we assess the effects of three successive years of exposure to combinations of elevated CO2 and O3 on growth responses in a five trembling aspen (Populus tremuloides) clonal mixture in a regenerating stand. The experiment is located in Rhinelander, Wisconsin, USA (45°N 89°W) and employs free air carbon dioxide and ozone enrichment (FACE) technology. The aspen stand was exposed to a factorial combination of four treatments consisting of elevated CO2 (560 ppm), elevated O3 (episodic exposure-90 μ1 1−1 hour−1), a combination of elevated CO2 and O3, and ambient control in 30 m treatment rings with three replications.