Light pollution affects invasive and native plant traits important to plant competition and herbivorous insects

Biological Invasions - Invasions in urban settings have been understudied in terms of how invasions are impacted by uniquely urban stressors, such as streetlights. Plant physiology and phenology...     

The mechanisms of plant photosynthetic acclimation to elevated CO2 concentration]

When measured at a same CO(2) concentration, net photosynthetic rate is often significantly lower in long-term high CO(2)-grown plants than the ambient CO(2)-grown ones. This phenomenon is termed photosynthetic acclimation or down-regulation. Although there have been many reports and reviews, the mechanism(s) of the photosynthetic acclimation is not very clear. Combining the work of the authors' group, this paper briefly reviews the progress in studies on the mechanism(s) of the photosynthetic acclimation to elevated CO(2). It is suggested that besides the possible effects of respiration enhancement and excessive photosynthate accumulation, RuBP carboxylation limitation and RuBP regeneration limitation are probably the main factors leading to the photosynthetic acclimation.     

Leaf quality and insect herbivory in model tropical plant communities after long-term exposure to elevated atmospheric CO2

Results from laboratory feeding experiments have shown that elevated atmospheric carbon dioxide can affect interactions between plants and insect herbivores, primarily through changes in leaf nutritional quality occurring at elevated CO₂. Very few data are available on insect herbivory in plant communities where insects can choose among species and positions in the canopy in which to feed. Our objectives were to determine the extent to which CO₂-induced changes in plant communities and leaf nutritional quality may affect herbivory at the level of the entire canopy. We introduced equivalent populations of fourth instar Spodoptera eridania, a lepidopteran generalist, to complex model ecosystems containing seven species of moist tropical plants maintained under low mineral nutrient supply. Larvae were allowed to feed freely for 14 days, by which time they had reached the seventh instar. Prior to larval introductions, plant communities had been continuously exposed to either 340 l CO₂ l^–1 or to 610 l CO₂ l^–1 for 1.5 years. No major shifts in leaf nutritional quality [concentrations of N, total non-structural carbohydrates (TNC), sugar, and starch; ratios of: C/N, TNC/N, sugar/N, starch/N; leaf toughness] were observed between CO₂ treatments for any of the species. Furthermore, no correlations were observed between these measures of leaf quality and leaf biomass consumption. Total leaf area and biomass of all plant communities were similar when caterpillars were introduced. However, leaf biomass of some species was slightly greater-and for other species slightly less (e.g. Cecropia peltata)-in communities exposed to elevated CO₂. Larvae showed the strongest preference for C. peltata leaves, the plant species that was least abundant in all communites, and fed relatively little on plants species which were more abundant. Thus, our results indicate that leaf tissue quality, as described by these parameters, is not necessarily affected by elevated CO₂ under relatively low nutrient conditions. Hence, the potential importance of CO₂-induced shifts in leaf nutritional quality, as determinants of herbivory, may be overestimated for many plant communities growing on nutrient-poor sites if estimates are based on traditional laboratory feeding studies. Finally, slight shifts in the abundance of leaf tissue of various species occurring under elevated CO₂ will probably not significantly affect herbivory by generalist insects. However, generalist insect herbivores appear to become more dependent on less-preferred plant species in cases where elevated CO₂ results in reduced availability of leaves of a favoured plant species, and this greater dependency may eventually affect insect populations adversely.     

大气CO_2浓度升高对植物-植食性昆虫的作用机制

大气二氧化碳浓度升高及其伴随的全球变暖引起国内外科学家的极大关注。CO2浓度升高主要通过改变植物的初级和次级代谢产物,影响以之为食的昆虫。本文结合作者近年来的研究成果,着重于以CO2浓度升高为作用因子,以植物和植食性昆虫的相互关系为对象,比较了咀嚼式口器昆虫与刺吸式口器昆虫对大气CO2浓度升高的响应特征,分析了不同取食类型昆虫-植物对大气CO2浓度升高的响应机制。     

Tree seedling growth in natural deep shade: functional traits related to interspecific variation in response to elevated CO2

The mechanisms for species-specific growth responses to changes in atmospheric CO₂ concentration within narrow ecological groups of species, such as shade-tolerant, late-successional trees, have rarely been addressed and are not well understood. In this study the underlying functional traits for interspecific variation in the biomass response to elevated CO₂ were explored for seedlings of five late-successional temperate forest tree species (Fagus sylvatica, Acer pseudoplatanus, Quercus robur, Taxus baccata, Abies alba). The seedlings were grown in the natural forest understorey in very low and low light microsites (an average of 1.3% and 3.4% full sun in this experiment), and were exposed to either current ambient CO₂ concentrations, 500, or 660 µl CO₂ l^-1 in 36 open-top chambers (OTC) over two growing seasons. Even across the narrow range of successional status and shade tolerance, the study species varied greatly in photosynthesis, light compensation point, leaf dark respiration (R_d), leaf nitrogen concentration, specific leaf area (SLA), leaf area ratio (LAR), and biomass allocation among different plant parts, and showed distinct responses to CO₂ in these traits. No single species combined all characteristics traditionally considered as adaptive to low light conditions. At very low light, the CO₂ stimulation of seedling biomass was related to increased LAR and decreased R_d, responses that were observed only in Fagus and Taxus. At slightly higher light levels, interspecific differences in the biomass response to elevated CO₂ were reversed and correlated best with leaf photosynthesis. The data provided here contribute to a mechanistic process-based understanding of distinct response patterns in co-occurring tree species to elevated CO₂ in natural deep shade. I conclude that the high variation in physiological and morphological traits among late-successional species, and the consequences for their responses to slight changes in resource availability, have previously been underestimated. The commonly used broad definitions of functional groups of species may not be sufficient for the understanding of recruitment success and dynamic changes in species composition of old-growth forests in response to rising concentrations of atmospheric CO₂.     

A link between plant diversity, elevated CO2 and soil nitrate

Interactive effects of reductions in plant species diversity and increases in atmospheric CO₂ were investigated in a long-term study in nutrient-poor calcareous grassland. Throughout the experiment, soil nitrate was persistently increased at low plant species diversity, and CO₂ enrichment reduced soil [NO₃^-] at all levels of plant species diversity. In our study, soil [NO₃^-] was unrelated to root length density, microbial biomass N, community legume contents, and experimental plant communities differed only little in total N pools. However, potential nitrification revealed exactly the same treatment effects as soil [NO₃^-], providing circumstantial evidence that nitrification rates drove the observed changes in [NO₃^-]. One possible explanation for plant diversity effects on nitrification lies in spatial and temporal interspecific differences in plant N uptake, which would more often allow accumulation of NH₄⁺ in part of the soil profile at low diversity than in more species-rich plant communities. Consequently, nitrification rates and soil [NO₃^-] would increase. Elevated CO₂ increased soil water contents, which may have improved NO₃^- diffusion to the root surface thereby reducing soil [NO₃^-]. Higher soil moisture at elevated CO₂ might also reduce nitrification rates due to less aerobic conditions. The accordance of the diversity effect on soil [NO₃^-] with previous experiments suggests that increased soil [NO₃^-] at low species diversity is a fairly general phenomenon, although the mechanisms causing high [NO₃^-] may vary. In contrast, experimental evidence for effects of CO₂ enrichment on soil [NO₃^-] is ambiguous, and the antagonistic interaction of plant species reductions and elevated CO₂ we have observed is thus probably less universal.     

Plant nutrient mobilization in temperate heathland responds to elevated CO2, temperature and drought

Temperate terrestrial ecosystems are currently exposed to increased atmospheric CO₂ and progressive climatic changes with increased temperature and periodical drought. We here present results from a field experiment, where the effects of these three main climate change related factors are investigated solely and in all combinations at a temperate heathland. Significant responses were found in the top soils below the two dominant species (Calluna vulgaris and Deschampsia flexuosa). During winter incubation, microbial immobilization of N and ammonification rate decreased in response to warming in Deschampsia soil, and microbial immobilization of N and P decreased in warmed Calluna soil. Warming tended to increase microbial N and P in Calluna but not in Deschampsia soil in fall, and more microbial C was accumulated under drought in Calluna soil. The effects of warming were often counteracted or erased when combined with CO₂ and drought. Below Deschampsia, the net nitrification rate decreased in response to drought and, while phosphorus availability and microbial P immobilization decreased, but nitrification increased in response to elevated CO₂. Furthermore, leaf litter decomposition of both species decreased in response to drought. These complex changes in availability and release of nutrients from soil organic matter turnover and mineralization in response to elevated CO₂ and climate change may influence the future plant carbon sequestration and species composition at temperate heathlands.     

Responses of an old-field plant community to interacting factors of elevated [CO2], warming, and soil moisture

Aims: The direct effects of atmospheric and climatic change factors—atmospheric[CO₂], air temperature and changes in precipitation—canshape plant community composition and alter ecosystem function.It is essential to understand how these factors interact tomake better predictions about how ecosystems may respond tochange. We investigated the direct and interactive effects of[CO₂], warming and altered soil moisture in open-top chambers(OTCs) enclosing a constructed old-field community to test howthese factors shape plant communities. Materials and methods: The experimental facility in Oak Ridge, TN, USA, made use of4-m diameter OTCs and rain shelters to manipulate [CO₂] (ambient,ambient + 300 ppm), air temperature (ambient, ambient + 3.5°C)and soil moisture (wet, dry). The plant communities within thechambers comprised seven common old-field species, includinggrasses, forbs and legumes. We tracked foliar cover for eachspecies and calculated community richness, evenness and diversityfrom 2003 to 2005. Important findings: This work resulted in three main findings: (1) warming had species-specificeffects on foliar cover that varied through time and were alteredby soil moisture treatments; (2) [CO₂] had little effect onindividual species or the community; (3) diversity, evennessand richness were influenced most by soil moisture, primarilyreflecting the response of one dominant species. We concludethat individualistic species responses to atmospheric and climaticchange can alter community composition and that plant populationsand communities should be considered as part of analyses ofterrestrial ecosystem response to climate change. However, predictionof plant community responses may be difficult given interactionsbetween factors and changes in response through time.     

Long-term acclimation to elevated pCO2 alters carbon metabolism and reduces growth in the Antarctic diatom Nitzschia lecointei

Increasing atmospheric CO₂ levels are driving changes in the seawater carbonate system, resulting in higher pCO₂ and reduced pH (ocean acidification). Many studies on marine organisms have focused on short-term physiological responses to increased pCO₂, and few on slow-growing polar organisms with a relative low adaptation potential. In order to recognize the consequences of climate change in biological systems, acclimation and adaptation to new environments are crucial to address. In this study, physiological responses to long-term acclimation (194 days, approx. 60 asexual generations) of three pCO₂ levels (280, 390 and 960 µatm) were investigated in the psychrophilic sea ice diatom Nitzschia lecointei. After 147 days, a small reduction in growth was detected at 960 µatm pCO₂. Previous short-term experiments have failed to detect altered growth in N. lecointei at high pCO₂, which illustrates the importance of experimental duration in studies of climate change. In addition, carbon metabolism was significantly affected by the long-term treatments, resulting in higher cellular release of dissolved organic carbon (DOC). In turn, the release of labile organic carbon stimulated bacterial productivity in this system. We conclude that long-term acclimation to ocean acidification is important for N. lecointei and that carbon overconsumption and DOC exudation may increase in a high-CO₂ world.     

Response of a Sphagnum bog plant community to elevated CO2 and N supply

The factors determining herbaceous canopy architecture are poorlyunderstood, especially in natural and semi-natural plant communities. Inthis study, we tested three main hypotheses: (1) the structure of herbaceouscanopies can be explained by the vertical distribution of functional groupsdefined by leaf width and the presence/absence of leaves on upright stem;(2) the degree of canopy stratification is greater in habitats that experiencelower spatial heterogeneity in the supply of light (i.e., grasslands as opposedto forest herb layers); and (3) there is significant variation among specieswithin a growth-form, with respect to their vertical position in thecanopy. We used plant foliage height distribution data from 14 grassland and 13forest herbaceous communities to test these hypotheses. A general linear mixedmodel was applied to specify the proportions of total variance in the foliageheight, accounted for by the fixed effects of plants' basicgrowth-form properties (growth-form) and community type(forest/grassland), and by the random effects of sampling site, samplingpoint, and individual species. We were also interested in the correlation ofthedegree of the stratification with various community characteristics(productivity, other canopy properties, species richness, variation ofspecies' traits) and light availability. There was some evidence ofoverall canopy stratification according toplant growth-form, since plants with leafy stem were locatedsignificantly higher. However, such a pattern of two more or less distinctlayers (grasses + upright forbs and rosette forbs) occurred withconsistency only in grasslands (greater homogeneity in light). Thebetween-species variation within a growth-form was a highlysignificant predictor of canopy vertical structure in the 27 communities. Theproportion of total observed variance, explainable throughspecies-specific effects, was comparable to that caused bybetween-site differences. The effect of community horizontal pattern wasless obvious, but still significant. The site by site analysis revealed thatthe degree to which horizontalpatchiness explained variation in vertical canopy structure was negativelyrelated to the relative importance of species-specific effects, showingthat small between-species differences lead to a more obviouswithin-community horizontal pattern, and vice versa. The upper bound ofthe degree of foliage stratification, according to growth-form, wasrelated to the variability of species light requirements and to relative (tocommunity pool size) richness, indicating that certain aspects of canopyarchitecture might be explained through community species composition anddiversity pattern.     

Genetic variation in plant below-ground response to elevated CO2 and two herbivore species

Aims

It is unclear how changing atmospheric conditions, including rising carbon dioxide concentration, influence interactions between above and below-ground systems and if intraspecific variation exists in this response.

Methods

We assessed interactive effects of atmospheric CO₂ concentration, above-ground herbivory, and plant genotype on root traits and mycorrhizal associations. Plants from five families of Asclepias syriaca, a perennial forb, were grown under ambient and elevated atmospheric CO₂ concentrations. Foliar herbivory by either lepidopteran caterpillars or phloem-feeding aphids was imposed. Mycorrhizal colonization, below-ground biomass, root biomass, and secondary defensive chemistry in roots were quantified.

Results

We observed substantial genetic variation among A. syriaca families in their mycorrhizal colonization levels in response to elevated CO₂ and herbivory treatments. Elevated CO₂ treatment increased root biomass in all genetic families, whereas foliar herbivory tended to decrease root biomass. Root cardenolide concentration and composition varied greatly among plant families, and elevated CO₂ treatment increased root cardenolides in two of the five plant families. Moreover, herbivores differentially affected the composition of cardenolides expressed below ground.

Conclusions

Increased atmospheric CO₂ has the potential to influence interactions among plants, herbivores and mycorrhizal fungi and intraspecific variation suggests that such interactions can evolve.     

The response of mycorrhizal colonization to elevated CO2 and climate change in Pascopyrum smithii and Bouteloua gracilis

Large intact soil cores of nearly pure stands of Pascopyrum smithii (western wheatgrass, C₃) and Bouteloua gracilis (blue grama, C₄) were extracted from the Central Plains Experimental Range in northeastern Colorado, USA and transferred to controlled environment chambers. Cores were exposed to a variety of water, temperature and CO₂ regimes for a total of four annual growth cycles. Root subsamples were harvested after the completion of the second and fourth growth cycles at a time corresponding to late winter, and were examined microscopically for the presence of mycorrhizae. After two growth cycles in the growth chambers, 54% of the root length was colonized in P. smithii, compared to 35% in blue grama. Field control plants had significantly lower colonization. Elevation of CO₂ increased mycorrhizal colonization in B. gracilis by 46% but had no effect in P. smithii. Temperatures 4° C higher than normal decreased colonization in P. smithii by 15%. Increased annual precipitation decreased colonization in both species. Simulated climate change conditions of elevated CO₂, elevated temperature and lowered precipitation decreased colonization in P. smithii but had less effect on B. gracilis. After four growth cycles in P. smithii, trends of treatments remained similar, but overall colonization rate decreased.     

Responses of fungal root colonization,plant cover and leaf nutrients to long‐term exposure to elevated atmospheric CO2 and warming in a subarctic birch forest understory

Responses of the mycorrhizal fungal community in terrestrial ecosystems to global change factors are not well understood. However, virtually all land plants form symbiotic associations with mycorrhizal fungi, with approximately 20% of the plants' net primary production transported down to the fungal symbionts. In this study, we investigated how ericoid mycorrhiza (ErM), fine endophytes (FE) and dark septate endophytes (DSE) in roots responded to elevated atmospheric CO₂ concentrations and warming in the dwarf shrub understory of a birch forest in the subarctic region of northern Sweden. To place the belowground results into an ecosystem context we also investigated how plant cover and nutrient concentrations in leaves responded to elevated atmospheric CO₂ concentrations and warming. The ErM colonization in ericaceous dwarf shrubs increased under elevated atmospheric CO₂ concentrations, but did not respond to warming following 6 years of treatment. This suggests that the higher ErM colonization under elevated CO₂ might be due to increased transport of carbon belowground to acquire limiting resources such as N, which was diluted in leaves of ericaceous plants under enhanced CO₂. The elevated CO₂ did not affect total plant cover but the plant cover was increased under warming, which might be due to increased N availability in soil. FE colonization in grass roots decreased under enhanced CO₂ and under warming, which might be due to increased root growth, to which the FE fungi could not keep up, resulting in proportionally lower colonization. However, no responses in aboveground cover of Deschampsia flexuosa were seen. DSE hyphal colonization in grass roots significantly increased under warmer conditions, but did not respond to elevated CO₂. This complex set of responses by mycorrhizal and other root‐associated fungi to global change factors of all the fungal types studied could have broad implications for plant community structure and biogeochemistry of subarctic ecosystems.     

A meta-analysis of elevated CO2 effects on woody plant mass, form, and physiology

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. Received: 16 May 1997 / Accepted: 9 September 1997     

Nitrogen and carbon partitioning in soybean under variable nitrogen supplies and acclimation to the prolonged action of elevated CO2

This study was conducted to determine reciprocal effects of low to high doses of nitrogenous fertilizer (N₃₀, N₄₀, N₅₀, N₆₀ and N₇₀ — 30, 40, 50, 60 and 70 kg ha⁻¹ respectively) and CO₂ enriched environment on C and N partitioning in soybean (Glycine max (L.) Merril cv JS-335). Plants were grown from seedling emergence to maturity inside open top chambers under ambient, AC (350±50 mol mol⁻¹) and elevated, EC (600±50 mol mol⁻¹) CO₂ and analyzed at seedling, vegetative, flowering, pod setting and maturity stages. Soybean responded to both CO₂ enrichment and N supply. Leaves, stem and root reserves at different growth stages were analyzed for total C and N contents. Consistent increase in the C contents of the leaf, stem and root was observed under EC than in AC. N contents in the different plant parts were found to be decreased under EC-grown plants specially at seedling and vegetative stage despite providing N doses to the soil. Significant increase observed for C to N dry mass ratio under EC in the root, stems and leaves at seedling and vegetative stage was decreased in the middle and later growth stages possibly due to combined impact of N doses to the soil and increased N₂ fixing activities due to EC conditions. Critical analysis of our findings reveals that the composition and partitioning of C and N of soybean under variable rates of N supply and CO₂ enrichment alter according to need under altered metabolic process. These changes eventually may lead to alteration in uptake of not only N but other essential nutrients also under changing atmosphere.     

The influence of plant species,fertilization and elevated CO2 on soil aggregate stability

We tested the effects of plant species, fertilization and elevated CO₂ on water-stable soil aggregation. Five annual grassland species and a plant community were grown in outdoor mesocosms for 4 years, with and without NPK fertilization, at ambient or elevated atmospheric CO₂ concentrations. Aggregate stability (resistance of aggregates to slaking) in the top 0.15 m of soil differed among plant species. However, the more diverse plant community did not enhance aggregate stability relative to most monocultures. Species differences in aggregate stability were positively correlated with soil active bacterial biomass, but did not correlate with root biomass or fungal length. Plant species did not affect aggregate stability lower in the soil profile (0.15–0.45 m), where soil biological activity is generally decreased. Elevated CO₂ and NPK fertilization altered many of the factors known to influence aggregation, but did not affect water-stable aggregation at either depth, in any of the plant treatments. These results suggest that global changes will alter soil structure primarily due to shifts in vegetation composition.     

Effects of elevated CO2 and plant genotype on interactions among cotton,aphids and parasitoids

Abstract Effects of CO₂ level (ambient vs. elevated) on the interactions among three cotton (Gossypium hirsutum) genotypes, the cotton aphid (Aphis gossypii Glover), and its hymenoptera parasitoid (Lysiphlebia japonica Ashrnead) were quantified. It was hypothesized that aphid‐parasitoid interactions in crop systems may be altered by elevated CO₂, and that the degree of change is influenced by plant genotype. The cotton genotypes had high (M9101), medium (HZ401) and low (ZMS13) gossypol contents, and the response to elevated CO₂ was genotype‐specific. Elevated CO₂ increased the ratio of total non‐structural carbohydrates to nitrogen (TNC: N) in the high‐gossypol genotype and the medium‐gossypol genotype. For all three genotypes, elevated CO₂ had no effect on concentrations of gossypol and condensed tannins. A. gossypii fitness declined when aphids were reared on the high‐gossypol genotype versus the low‐gossypol genotype under elevated CO₂. Furthermore, elevated CO₂ decreased the developmental time of L. japonica associated with the high‐gossypol genotype and the low‐gossypol genotype, but did not affect parasitism or emergence rates. Our study suggests that the abundance of A. gossypii on cotton will not be directly affected by increases in atmospheric CO₂. We speculate that A. gossypii may diminish in pest status in elevated CO₂ and high‐gossypol genotype environments because of reduced fitness to the high‐gossypol genotype and shorter developmental time of L. japonica.     

Root system adjustments: regulation of plant nutrient uptake and growth responses to elevated CO2

Nutrients such as nitrogen (N) and phosphorus (P) often limit plant growth rate and production in natural and agricultural ecosystems. Limited availability of these nutrients is also a major factor influencing long-term plant and ecosystem responses to rising atmospheric CO₂ levels, i.e., the commonly observed short-term increase in plant biomass may not be sustained over the long-term. Therefore, it is critical to obtain a mechanistic understanding of whether elevated CO₂ can elicit compensatory adjustments such that acquisition capacity for minerals increases in concert with carbon (C) uptake. Compensatory adjustments such as increases in (a) root mycorrhizal infection, (b) root-to-shoot ratio and changes in root morphology and architecture, (c) root nutrient absorption capacity, and (d) nutrient-use efficiency can enable plants to meet an increased nutrient demand under high CO₂. Here we examine the literature to assess the extent to which these mechanisms have been shown to respond to high CO₂. The literature survey reveals no consistent pattern either in direction or magnitude of responses of these mechanisms to high CO₂. This apparent lack of a pattern may represent variations in experimental protocol and/or interspecific differences. We found that in addressing nutrient uptake responses to high CO₂ most investigators have examined these mechanisms in isolation. Because such mechanisms can potentially counterbalance one another, a more reliable prediction of elevated CO₂ responses requires experimental designs that integrate all mechanisms simultaneously. Finally, we present a functional balance (FB) model as an example of how root system adjustments and nitrogen-use efficiency can be integrated to assess growth responses to high CO₂. The FB model suggests that the mechanisms of increased N uptake highlighted here have different weights in determining overall plant responses to high CO₂. For example, while changes in root-to-shoot biomass allocation, r, have a small effect on growth, adjustments in uptake rate per unit root mass, [`(n)]\bar \nu , and photosynthetic N use efficiency, p*, have a significantly greater leverage on growth responses to elevated CO₂ except when relative growth rate (RGR) reaches its developmental limit, maximum RGR (RGR_max).     

Antioxidants in sun and shade leaves of sour orange trees (Citrus aurantium) after long-term acclimation to elevated CO2

Antioxidative systems and the contents of pigments, malondialdehyde,soluble protein, and carbohydrate were investigated in sun-and shade-acclimated leaves of sour orange (Citrus aurantium)trees that had been grown for 7.5 years under ambient and elevated(+300 µmol mol^–1) atmospheric CO₂ concentrations.Sunacclimated leaves contained higher ascorbate, glutathioneand soluble carbohydrate contents and higher catalase activitiesthan shade-acclimated leaves. The activities of superoxide dismutases,which belonged to the family of Cu/Zn-isozymes, were similarin sunand shade-acclimated leaves and decreased in responseto enhanced CO₂. In shade-acclimated leaves, none of the otherparameters studied was affected by elevated CO₂. In sun-acclimatedleaves elevated CO₂ caused increases in carbohydrate and ascorbatecontents. There was no evidence for enhanced lipid peroxidationas assessed from the determination of the malondialdehyde contentsunder either conditions. Key words: Ascorbate, catalase, CO₂ enrichment, global change, glutathione, superoxide dismutase