Genotypic variation in the response of two perennial grass species to elevated carbon dioxide

Shoot and reproductive biomass of genotypes of Bromus erectus and Dactylis glomerata grown in competition at ambient and elevated CO₂ were examined for 2 consecutive years in order to test whether genetic variation in those traits exists and whether it is maintained over time. At the species level, a positive CO₂ response of shoot biomass of both species was only found in the first year of treatment. At the genotype level, no significant CO₂2genotype interaction was found at any single harvest either for vegetative or reproductive biomass of either species. Analysis over time, however, indicated that there is a potential for evolutionary adaptation only for D. glomerata: (1) repeated measures ANOVA detected a marginally significant CO₂2genotype2time interaction for shoot biomass, because the range of the genotypes CO₂ response increased over time; (2) genotypes that displayed the highest response during the first year under elevated CO₂ also showed the highest response the second year. Null (B. erectus) or weak (D. glomerata) selective potentials of elevated CO₂ were detected in this experiment, but short time series could underestimate this potential with perennial species.     

Modeling dynamic understory photosynthesis of contrasting species in ambient and elevated carbon dioxide

Dynamic responses of understory plants to sunflecks have been extensively studied, but how much differences in dynamic light responses affect daily photosynthesis (A_day) is still the subject of active research. Recent models of dynamic photosynthesis have provided a quantitative tool that allows the critical assessment of the importance of these sunfleck responses on A_day. Here we used a dynamic photosynthesis model to assess differences in four species that were growing in ambient and elevated CO₂. We hypothesized that Liriodendron tulipifera, a species with rapid photosynthetic induction gain and slow induction loss, would have the least limitations to sunfleck photosynthesis relative to the other three species (Acer rubrum, Cornus florida, Liquidambar styraciflua). As a consequence, L. tulipifera should have the highest A_day in an understory environment, despite being the least shade tolerant of the species tested. We further hypothesized that daily photosynthetic enhancement by elevated CO₂ would differ from enhancement levels observed during light-saturated, steady-state measurements. Both hypotheses were supported by the model results under conditions of low daily photosynthetic photon flux density (PFD; <3% of the above-canopy PFD). However, under moderate PFD (10-20% of the above-canopy PFD), differences in dynamic sunfleck responses had no direct impact on A_day for any of the species, since stomatal and photosynthetic induction limitations to sunfleck photosynthesis were small. Thus, the relative species ranking in A_day under moderate PFD closely matched their rankings in steady-state measurements of light-saturated photosynthesis. Similarly, under elevated CO₂, enhancement of modeled A_day over A_day at ambient CO₂ matched the enhancement measured under light saturation. Thus, the effects of species-specific differences in dynamic sunfleck responses, and differences in elevated CO₂ responses of daily photosynthesis, are most important in marginal light environments.     

Differential impacts of long-term (CO<Subscript>2</Subscript>) and O<Subscript>3</Subscript> exposure on growth of northern conifer and deciduous tree species

The long-term effects of elevated CO₂ and CO₂+O₃ concentrations on the growth allocation in northern provenances of Norway spruce [Picea abies (L.) Karst.], Scots pine [Pinus sylvestris (L.)] and pubescent birch clones (Betula pubescens Ehrh.) were examined in open-top chambers after a 4-year-long experiment. The total biomass responses of the tree seedlings to increased CO₂ and CO₂+O₃ concentrations were not statistically significant and varied between the provenances and species. The seedlings of northern origin were the least sensitive in their response to treatments. The total biomass of the Norway spruce seedlings slightly decreased in response to CO₂ in three provenances. Scots pine from the local provenance had a slight biomass increase after elevated CO₂+O₃ treatment. The slower-growing birch clone seemed to benefit from elevated CO₂, whereas in the faster-growing clone, reductions in biomass accumulation were seen. The combined CO₂+O₃ treatment reduced the positive effects of elevated CO₂, especially in the slower-growing birches. Observations of significant effects were limited to a few parameters. Carbon dioxide treatment decreased needle dry weight of Norway spruce in one northern provenance. The needle and wood dry weight increased (CO₂ + O₃) in local Scots pine. Significant birch response was limited to increased fine root density (O₃ + CO₂) in the inland clone. The diverse effects of elevated CO₂ and CO₂ +O₃ on seedling growth and biomass provide evidence that exposure of northern trees to the enhanced variable CO₂ and O₃ concentrations of the future will have varied effects on the growth of these species. The direction and magnitude of those effects will differ depending on species and origins.     

Four-year growth dynamics of beech-spruce model ecosystems under CO2 enrichment on two different forest soils

To elucidate how atmospheric CO₂ enrichment, enhanced nutrient supply and soil quality interact to affect regrowth of temperate forests, young Fagus sylvatica and Picea abies trees were grown together in large model ecosystems. Identical communities were established on a nutrient-poor acidic and on a more fertile calcareous soil and tree growth, leaf area index, fine root density and soil respiration monitored over four complete growing seasons. Biomass responses to CO₂ enrichment and enhanced N supply at the end of the experiment reflected compound interest effects of growth stimulation during the first two to three seasons rather than persistent stimulation over the whole duration of the experiment. Whereas biomass of Picea was enhanced in elevated CO₂ on both soils, Fagus responded negatively to CO₂ on acidic but positively on calcareous soil. Biomass of both species profited from enhanced N supply on the poor acidic soil only. Leaf area index on both soils was greater in high N supply as a consequence of a stimulation early in the experiment, but was unaffected by CO₂ enrichment. Fine root density on acidic soil was increased in high N supply, but this did not stimulate soil respiration rate. In contrast, elevated CO₂ stimulated both fine root density and soil CO₂ efflux on calcareous soil, especially towards the end of the experiment. Our experiment suggests that future species dominance in beech-spruce forests is likely to change in response to CO₂ enrichment, but this response is subject to complex interactions with environmental factors other than CO₂, particularly soil type.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Diverse Responses of Maple Saplings to Forest Light Regimes

Seedlings of 11 species of forest maples (AcerL.) were grownoutdoors from budburst to senescence under three light regimes:‘gap centre under clear skies’ (approx. 20% opensky irradiance; red:far-red ratio=1.12); ‘gap centre undercloudy skies’ (1.5%, ratio=1.03); and ‘gap edge’(2.5%, ratio=0.6). Seedlings grown under the gap centre (clearsky) regime had significantly greater height growth, greaterspecific leaf mass, higher root:shoot ratio, greater investmentin roots, higher leaf nitrogen concentrations, greater chlorophylla:bratio,lower photosynthetic rates under dim light, higher maximum photosyntheticrate, higher stomatal conductance, and lower leaf internal CO₂concentrationscompared with those grown in either gap edge or gap centre (cloudy)regimes. Responses to the gap edgevs.gap centre (cloudy) treatmentsdiffer little, suggesting that shade acclimation in forest mapleseedlings is mainly a response to light intensity rather thanspectral quality. The ubiquitous and, except for leaf internalCO₂concentration, highly significant interspecific variationin traits was broad-ranging and continuous. These results suggestthat (1) the responses to light quality found in shade intolerantherbaceous and woody species growing in more open habitats maynot have a selective advantage in seedlings of shade tolerantforest trees, and (2) the adaptive plastic response to understoreyvs.gapenvironments in forest maples, which is qualitatively consistentacross species, is founded on co-ordinated, small shifts insets of functionally inter-related traits.Copyright 1998 Annalsof Botany Company Acer,forest gap heterogeneity, plasticity, specific leaf mass, photosynthesis, leaf chlorophyll, nitrogen, stomatal density, root growth, root:shoot ratio, growth form.     

Responses of communities of tropical tree species to elevated CO2 in a forest clearing

Communities of ten species of tropical forest tree seedlings from three successional classes were grown at ambient and elevated CO₂ in large open-top chambers on the edge of a forest in Panamá. Communities grew from 20?cm to approximately 2?m in height in 6 months. No enhancements in plant biomass accumulation occurred under elevated CO₂ either in the whole communities or in growth of individual species. Reductions in leaf area index under elevated CO₂ were observed, as were decreases in leaf nitrogen concentrations and increases in the C:N ratio of leaf tissue. Species tended to respond individualistically to elevated CO₂, but some generalizations of how successional groupings responded could be made. Early and mid-successional species generally showed greater responses to elevated CO₂ than late-successional species, particularly with respect to increases in photosynthetic rates and leaf starch concentrations, and reductions in leaf area ratio. Late-successional species showed greater increases in C:N ratios in response to elevated CO₂ than did other species. Our results indicate that there may not be an increase in the growth of regenerating tropical forest under elevated CO₂, but that there could be changes in soil nutrient availability because of reductions in leaf tissue quality, particularly in late-successional species.     

Consequences of CO2 and light interactions for leaf phenology, growth, and senescence in Quercus rubra

We investigated how light and CO₂ levels interact to influence growth, phenology, and the physiological processes involved in leaf senescence in red oak (Quercus rubra) seedlings. We grew plants in high and low light and in elevated and ambient CO₂. At the end of three years of growth, shade plants showed greater biomass enhancement under elevated CO₂ than sun plants. We attribute this difference to an increase in leaf area ratio (LAR) in shade plants relative to sun plants, as well as to an ontogenetic effect: as plants increased in size, the LAR declined concomitant with a decline in biomass enhancement under elevated CO₂ Elevated CO₂ prolonged the carbon gain capacity of shade‐grown plants during autumnal senescence, thus increasing their functional leaf lifespan. The prolongation of carbon assimilation, however, did not account for the increased growth enhancement in shade plants under elevated CO₂. Elevated CO₂ did not significantly alter leaf phenology. Nitrogen concentrations in both green and senesced leaves were lower under elevated CO₂ and declined more rapidly in sun leaves than in shade leaves. Similar to nitrogen concentration, the initial slope of A/C_i curves indicated that Rubisco activity declined more rapidly in sun plants than in shade plants, particularly under elevated CO₂. Absolute levels of chlorophyll were affected by the interaction of CO₂ and light, and chlorophyll content declined to a minimal level in sun plants sooner than in shade plants. These declines in N concentration, in the initial slope of A/C_i curves, and in chlorophyll content were consistent with declining photosynthesis, such that elevated CO₂ accelerated senescence in sun plants and prolonged leaf function in shade plants. These results have implications for the carbon economy of seedlings and the regeneration of red oak under global change conditions.     

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).     

Effects of elevated CO<Subscript>2</Subscript> on foliar chemistry of saplings of nine species of tropical tree

This study examined the effects of elevated CO₂ on secondary metabolites for saplings of tropical trees. In the first experiment, nine species of trees were grown in the ground in open-top chambers in central Panama at ambient and elevated CO₂ (about twice ambient). On average, leaf phenolic contents were 48% higher under elevated CO₂. Biomass accumulation was not affected by CO₂, but starch, total non-structural carbohydrates and C/N ratios all increased. In a second experiment with Ficus, an early successional species, and Virola, a late successional species, treatments were enriched for both CO₂ and nutrients. For both species, nutrient fertilization increased plant growth and decreased leaf carbohydrates, C/N ratios and phenolic contents, as predicted by the carbon/nutrient balance hypothesis. Changes in leaf C/N levels were correlated with changes in phenolic contents for Virola (r=0.95, P<0.05), but not for Ficus. Thus, elevated CO₂, particularly under conditions of low soil fertility, significantly increased phenolic content as well as the C/N ratio of leaves. The magnitude of the changes is sufficient to negatively affect herbivore growth, survival and fecundity, which should have impacts on plant/herbivore interactions.     

Biomass responses to intraspecific competition differ between wild species and CO2

Aims: How growth of wild and crop species responds to global environmentalperturbations has both ecological and agricultural significancein a changing world. The primary aim of this synthesis was toquantitatively assess the interactive effects of intraspecificcompetition and elevated CO₂ on biomass production in herbaceousspecies. Methods: Using meta-analytical techniques, we synthesized data from publicationsbefore 2006 that reported biomass responses to elevated CO₂in 321 herbaceous species grown in isolation or in competitionwith con-specific individuals. Important findings: Intraspecific competition differentially modified biomass responsesto elevated CO₂ in wild and crop species. For example, competitionreduced CO₂ stimulation of total biomass (W_T) from 27 to 23%in wild species, but by a much greater magnitude, i.e., from43 to 32% in crops. Competition had no effect on responses ofeither above- (W_AG) or below-ground (W_BG) biomass to elevatedCO₂ in wild species, but significantly diminished CO₂ enhancementof W_AG, although not of W_BG, in crops. Considerable variationswere found among functional groups in the modification of growthresponses to elevated CO₂ by intraspecific competition, whichexerted greater depression on CO₂ enhancement in C₃ than inC₄ species and in non-legumes than in legumes. Elevated CO₂affected leaf and stem growth of individually grown C₃ graminoidsand forbs similarly, but increased leaf growth only in C₄ graminoidsand stem growth only in C₄ forbs. Results from this synthesisdemonstrated that intraspecific competition differentially affectedgrowth responses to elevated CO₂ in wild and crop species. Thewild–crop species differences will have important implicationsfor understanding primary production by herbaceous species inboth natural and agricultural ecosystems in the future whenatmospheric CO₂ is significantly higher than the current level.     

Photosynthesis in an invasive grass and native forb at elevated CO2 during an El Niño year in the Mojave Desert

Annual and short-lived perennial plant performance during wet years is important for long-term persistence in the Mojave Desert. Additionally, the effects of elevated CO₂ on desert plants may be relatively greater during years of high resource availability compared to dry years. Therefore, during an El Niño year at the Nevada Desert FACE Facility (a whole-ecosystem CO₂ manipulation), we characterized photosynthetic investment (by assimilation rate-internal CO₂ concentration relationships) and evaluated the seasonal pattern of net photosynthesis (A_net) and stomatal conductance (g_s) for an invasive annual grass, Bromus madritensis ssp. rubens and a native herbaceous perennial, Eriogonum inflatum. Prior to and following flowering, Bromus showed consistent increases in both the maximum rate of carboxylation by Rubisco (V_Cmax) and the light-saturated rate of electron flow (J_max) at elevated CO₂. This resulted in greater A_net at elevated CO₂ throughout most of the life cycle and a decrease in the seasonal decline of maximum midday A_net upon flowering as compared to ambient CO₂. Eriogonum showed significant photosynthetic down-regulation to elevated CO₂ late in the season, but the overall pattern of maximum midday A_net was not altered with respect to phenology. For Eriogonum, this resulted in similar levels of A_net on a leaf area basis as the season progressed between CO₂ treatments, but greater photosynthetic activity over a typical diurnal period. While g_s did not consistently vary with CO₂ in Bromus, it did decrease in Eriogonum at elevated CO₂ throughout much of the season. Since the biomass of both plants increased significantly at elevated CO₂, these patterns of gas exchange highlight the differential mechanisms for increased plant growth. The species-specific interaction between CO₂ and phenology in different growth forms suggests that important plant strategies may be altered by elevated CO₂ in natural settings. These results indicate the importance of evaluating the effects of elevated CO₂ at all life cycle stages to better understand the effects of elevated CO₂ on whole-plant performance in natural ecosystems.     

Microbial community composition and function beneath temperate trees exposed to elevated atmospheric carbon dioxide and ozone

We hypothesized that changes in plant growth resulting from atmospheric CO₂ and O₃ enrichment would alter the flow of C through soil food webs and that this effect would vary with tree species. To test this idea, we traced the course of C through the soil microbial community using soils from the free-air CO₂ and O₃ enrichment site in Rhinelander, Wisconsin. We added either ¹³C-labeled cellobiose or ¹³C-labeled N-acetylglucosamine to soils collected beneath ecologically distinct temperate trees exposed for 3 years to factorial CO₂ (ambient and 200 µl l^-1 above ambient) and O₃ (ambient and 20 µl l^-1 above ambient) treatments. For both labeled substrates, recovery of ¹³C in microbial respiration increased beneath plants grown under elevated CO₂ by 29% compared to ambient; elevated O₃ eliminated this effect. Production of ¹³C-CO₂ from soils beneath aspen (Populus tremuloides Michx.) and aspen-birch (Betula papyrifera Marsh.) was greater than that beneath aspen-maple (Acer saccharum Marsh.). Phospholipid fatty acid analyses (¹³C-PLFAs) indicated that the microbial community beneath plants exposed to elevated CO₂ metabolized more ¹³C-cellobiose, compared to the microbial community beneath plants exposed to the ambient condition. Recovery of ¹³C in PLFAs was an order of magnitude greater for N-acetylglucosamine-amended soil compared to cellobiose-amended soil, indicating that substrate type influenced microbial metabolism and soil C cycling. We found that elevated CO₂ increased fungal activity and microbial metabolism of cellobiose, and that microbial processes under early-successional aspen and birch species were more strongly affected by CO₂ and O₃ enrichment than those under late-successional maple.     

The effects of elevated CO2 on seed production and seedling recruitment in a sheep-grazed pasture

Seed production and seedling recruitment were measured over 2 years under ambient (360 ppm) and elevated (475 ppm) atmospheric CO₂ in a free air carbon dioxide enrichment (FACE) experiment, carried out in a sheep-grazed pasture on dry, sandy soil in New Zealand. In both years elevated CO₂ led to more dispersed seeds of the grasses Anthoxanthum odoratum, Lolium perenne and Poa pratensis, the legumes Trifolium repens and T. subterraneum and the herbs Hypochaeris radicata and Leontodon saxatilis. The increased seed dispersal in A. odoratum, H. radicata, Leontodon saxatilis and T. repens reflected both more inflorescences per unit area and more seeds per inflorescence under elevated CO₂. The increased seed dispersal in Lolium perenne, P. pratensis and T. subterraneum was due solely to more inflorescences per unit area. The number of seedlings that emerged and survived to at least 7 months of age was increased by elevated CO₂ for H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in both years and for A. odoratum and Lolium perenne in the first year. For species where increased seedling recruitment was noted, there was a significant positive correlation between seed production in summer and seedling emergence in the following autumn and winter, and sowing 200 extra seeds per species m^-2 resulted in more seedlings compared to unsown controls. Elevated CO₂ did not affect seedling survival in any species. There was no measurable effect of elevated CO₂ on canopy and soil surface conditions or soil moisture at the time of seedling emergence. The results suggest the dominant effect of elevated CO₂ on seedling recruitment in this pasture was an indirect one, reflecting effects on the number of seeds produced. The biomass of H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in the above-ground vegetation was greater under elevated than ambient CO₂. However, the size of individual seedlings and mature plants of these four species was unaffected by elevated CO₂. The results indicate an important way elevated CO₂ influenced plant species composition in this pasture was through changes in the pattern of seedling recruitment.     

Elevated CO2 influences nutrient availability in young beech-spruce communities on two soil types

The objectives of this study were to investigate how different soil types and elevated N deposition (0.7 vs 7 g N m^-2a^-1) influence the effects of elevated CO₂ (370 vs 570 µmol CO₂ mol^-1) on soil nutrients and net accumulation of N, P, K, S, Ca, Mg, Fe, Mn, and Zn in spruce (Picea abies) and beech (Fagus sylvatica). Model ecosystems were established in large open-top chambers on two different forest soils: a nutrient-poor acidic loam and a nutrient-rich calcareous sand. The response of net nutrient accumulation to elevated atmospheric CO₂ depended upon soil type (interaction soil 2 CO₂, P<0.05 for N, P, K, S, Ca, Mg, Zn) and differed between spruce and beech. On the acidic loam, CO₂ enrichment suppressed net accumulation of all nutrients in beech (P<0.05 for P, S, Zn), but stimulated it for spruce (P<0.05 for Fe, Zn) On the nutrient-rich calcareous sand, increased atmospheric CO₂ enhanced nutrient accumulation in both species significantly. Increasing the N deposition did not influence the CO₂ effects on net nutrient accumulation with either soil. Under elevated atmospheric CO₂, the accumulation of N declined relative to other nutrients, as indicated by decreasing ratios of N to other nutrients in tree biomass (all ratios: P<0.001, except the N to S ratio). In both the soil and soil solution, elevated CO₂ did not influence concentrations of base cations and available P. Under CO₂ enrichment, concentrations of exchangeable NH₄⁺ decreased by 22% in the acidic loam and increased by 50% in the calcareous sand (soil 2 CO₂, P<0.001). NO₃^- concentrations decreased by 10-70% at elevated CO₂ in both soils (P<0.01).     

Feeding rates of the woodlouse Armadillidium vulgare on herb litters produced at two levels of atmospheric CO2

The consumption and assimilation rates of the woodlouse Armadillidium vulgare were measured on leaf litters from five herb species grown and naturally senesced at 350 and 700 µl l^-1 CO₂. Each type of litter was tested separately after 12, 30 and 45 days of decomposition at 18°C. The effects of elevated CO₂ differed depending on the plant species. In Medicago minima (Fabaceae), the CO₂ treatment had no significant effect on consumption and assimilation. In Tyrimnus leucographus (Asteraceae), the CO₂ treatment had no significant effect on consumption, but the elevated CO₂ litter was assimilated at a lower rate than the ambient CO₂ litter after 30 days of decomposition. In the three other species, Galactites tomentosa (Asteraceae), Trifolium angustifolium (Fabaceae) and Lolium rigidum (Poaceae), the elevated CO₂ litter was consumed and/or assimilated at a higher rate than the ambient CO₂ litter. Examination of the nitrogen contents in these three species of litter did not support the hypothesis of compensatory feeding, i.e. an increase in woodlouse consumption to compensate for low nitrogen content of the food. Rather, the results suggest that in herbs that were unpalatable at the start of the experiment (Galactites, Trifolium and Lolium), more of the the litter produced at 700 µl l^-1 CO₂ was consumed than of that produced at 350 µl l^-1 because inhibitory factors were eliminated faster during decomposition.     

Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees

Among 13 tropical tree species on Barro Colorado Island, species with high seedling mortality rates during the first year in shade had higher reltive growth rates (RGR) from germination to 2 months in both sun (23% full sun) and shade [2%, with and without lowered red: far red (R:FR) ratio] than shade tolerant species. Species with higher RGR in sun also had higher RGR in shade. These interspecific trends could be explained by differences in morphological traits and allocation paterns among species. Within each light regime, seedlings of shade-intolerant species had lower root: shoot ratios, higher leaf mass per unit area, and higher leaf area ratios (LAR) than shade tolerant species. In contrast, leaf gas exchange characteristics, or acclimation potential in these traits, had no relationship with seedling mortality rates in shade. In both shade tolerant and intolerant species, light saturated photosynthesis rates, dark respiration, and light compensation points were higher for sungrown seedlings than for shade-grown seedlings. Differences in R:FR ratio in shade did not affect gas exchange, allocation patterns, or growth rates of any species. Survival of young tree seedlings in shade did not depend on higher net photosynthesis or biomass accumulation rates in shade. Rather, species with higher RGR died faster in shade than species with lower RGR. This trend could be explained if survival depends on morphological characteristics likely to enhance defense against herbivores and pathogens, such as dense and tough leaves, a well-established root system, and high wood density. High construction costs for these traits, and low LAR as a consequence of these traits, should result in lower rates of whole-plant carbon gain and RGR for shade tolerant species than shade-intolerant species in shade as well as in sun.     

The effect of elevated carbon dioxide and ozone on leaf- and branch-level photosynthesis and potential plant-level carbon gain in aspen

Two aspen (Populus tremuloides Michx.) clones, differing in O₃ tolerance, were grown in a free-air CO₂ enrichment (FACE) facility near Rhinelander, Wisconsin, and exposed to ambient air, elevated CO₂, elevated O₃ and elevated CO₂+O₃. Leaf instantaneous light-saturated photosynthesis (P_S) and leaf areas (A) were measured for all leaves of the current terminal, upper (current year) and the current-year increment of lower (1-year-old) lateral branches. An average, representative branch was chosen from each branch class. In addition, the average photosynthetic rate was estimated for the short-shoot leaves. A summing approach was used to estimate potential whole-plant C gain. The results of this method indicated that treatment differences were more pronounced at the plant- than at the leaf- or branch-level, because minor effects within modules accrued in scaling to plant level. The whole-plant response in C gain was determined by the counteracting changes in P_S and A. For example, in the O₃-sensitive clone (259), inhibition of P_S in elevated O₃ (at both ambient and elevated CO₂) was partially ameliorated by an increase in total A. For the O₃-tolerant clone (216), on the other hand, stimulation of photosynthetic rates in elevated CO₂ was nullified by decreased total A.     

Photosynthesis, photoinhibition, and nitrogen use efficiency in native and invasive tree ferns in Hawaii

Photosynthetic gas exchange, chlorophyll fluorescence, nitrogen use efficiency, and related leaf traits of native Hawaiian tree ferns in the genus Cibotium were compared with those of the invasive Australian tree fern Sphaeropteris cooperi in an attempt to explain the higher growth rates of S. cooperi in Hawaii. Comparisons were made between mature sporophytes growing in the sun (gap or forest edge) and in shady understories at four sites at three different elevations. The invasive tree fern had 12-13 cm greater height increase per year and approximately 5 times larger total leaf surface area per plant compared to the native tree ferns. The maximum rates of photosynthesis of S. cooperi in the sun and shade were significantly higher than those of the native Cibotium spp (for example, 11.2 and 7.1 µmol m^-2 s^-1, and 5.8 and 3.6 µmol m^-2 s^-1 respectively for the invasive and natives at low elevation). The instantaneous photosynthetic nitrogen use efficiency of the invasive tree fern was significantly higher than that of the native tree ferns, but when integrated over the life span of the frond the differences were not significant. The fronds of the invasive tree fern species had a significantly shorter life span than the native tree ferns (approximately 6 months and 12 months, respectively), and significantly higher nitrogen content per unit leaf mass. The native tree ferns growing in both sun and shade exhibited greater photoinhibition than the invasive tree fern after being experimentally subjected to high light levels. The native tree ferns recovered only 78% of their dark-acclimated quantum yield (F_v/F_m), while the invasive tree fern recovered 90% and 86% of its dark-acclimated F_v/F_m when growing in sun and shade, respectively. Overall, the invasive tree fern appears to be more efficient at capturing and utilizing light than the native Cibotium species, particularly in high-light environments such as those associated with high levels of disturbance.     

Effects of Elevated Carbon Dioxide on Plant Biomass Production and Competition in a Simulated Neutral Grassland Community

Using open-top chambers, four prominent species (Lolium perenne,Cynosurus cristatus, Holcus lanatusandAgrostis capillaris) ofIrish neutral grasslands were grown at ambient and elevated(700 µmol mol^-1) atmospheric CO₂for a period of 8 months.The effects of interspecific competition on plant responsesto CO₂enrichment were investigated by growing the species ina four-species mixture. The results indicate that the speciesdiffer in their ability to respond to elevated CO₂. CO₂-enrichmenthad the largest effect on the biomass production ofH. lanatus,but substantial stimulations in biomass production were alsofound for the other three species. The CO₂-stimulation of biomassproduction forH. lanatuswas accompanied by increased tillering.In addition, reductions in specific leaf area were found forall species. Exposure to elevated CO₂increased the communitybiomass of the four-species mixture. This increase can be mainlyattributed to a significant increase in the biomass ofH. lanatusatelevated CO₂. No statistically-significant changes in speciescomposition of community biomass were found. However,H. lanatusdidincrease its share of community biomass at each of the harvests,with the other three species, mainlyL. perenne, suffering lossesin their shares at elevated CO₂. The results show that: (1)the species varied in their response to elevated CO₂; and (2)species composition in natural plant communities is likely tochange at elevated CO₂, but these changes may occur rather slowly.Much longer periods of exposure to elevated atmospheric CO₂maybe required to permit detection of significant changes in speciescomposition.Copyright 1998 Annals of Botany Company Carbon dioxide (CO₂) enrichment, competition, Lolium perenne,Cynosurus cristatus, Holcus lanatus, Agrostis capillaris, biomass, specific leaf area, tillering.