An alpine treeline in a carbon dioxide-rich world: synthesis of a nine-year free-air carbon dioxide enrichment study

We evaluated the impacts of elevated CO₂ in a treeline ecosystem in the Swiss Alps in a 9-year free-air CO₂ enrichment (FACE) study. We present new data and synthesize plant and soil results from the entire experimental period. Light-saturated photosynthesis (A _max) of ca. 35-year-old Larix decidua and Pinus uncinata was stimulated by elevated CO₂ throughout the experiment. Slight down-regulation of photosynthesis in Pinus was consistent with starch accumulation in needle tissue. Above-ground growth responses differed between tree species, with a 33 % mean annual stimulation in Larix but no response in Pinus. Species-specific CO₂ responses also occurred for abundant dwarf shrub species in the understorey, where Vaccinium myrtillus showed a sustained shoot growth enhancement (+11 %) that was not apparent for Vaccinium gaultherioides or Empetrum hermaphroditum. Below ground, CO₂ enrichment did not stimulate fine root or mycorrhizal mycelium growth, but increased CO₂ effluxes from the soil (+24 %) indicated that enhanced C assimilation was partially offset by greater respiratory losses. The dissolved organic C (DOC) concentration in soil solutions was consistently higher under elevated CO₂ (+14 %), suggesting accelerated soil organic matter turnover. CO₂ enrichment hardly affected the C–N balance in plants and soil, with unaltered soil total or mineral N concentrations and little impact on plant leaf N concentration or the stable N isotope ratio. Sustained differences in plant species growth responses suggest future shifts in species composition with atmospheric change. Consistently increased C fixation, soil respiration and DOC production over 9 years of CO₂ enrichment provide clear evidence for accelerated C cycling with no apparent consequences on the N cycle in this treeline ecosystem.     

Wood properties and ring width responses to long-term atmospheric CO2 enrichment in field-grown loblolly pine (Pinus taeda L.)

Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO₂ partial pressures (pCO₂) and with a CO₂-enriched atmosphere (+ 30 Pa pCO₂ compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO₂. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO₂-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO₂ enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.     

Soil warming and CO2 enrichment induce biomass shifts in alpine tree line vegetation

Responses of alpine tree line ecosystems to increasing atmospheric CO₂ concentrations and global warming are poorly understood. We used an experiment at the Swiss tree line to investigate changes in vegetation biomass after 9 years of free air CO₂ enrichment (+200 ppm; 2001–2009) and 6 years of soil warming (+4 °C; 2007–2012). The study contained two key tree line species, Larix decidua and Pinus uncinata, both approximately 40 years old, growing in heath vegetation dominated by dwarf shrubs. In 2012, we harvested and measured biomass of all trees (including root systems), above‐ground understorey vegetation and fine roots. Overall, soil warming had clearer effects on plant biomass than CO₂ enrichment, and there were no interactive effects between treatments. Total plant biomass increased in warmed plots containing Pinus but not in those with Larix. This response was driven by changes in tree mass (+50%), which contributed an average of 84% (5.7 kg m^?2) of total plant mass. Pinus coarse root mass was especially enhanced by warming (+100%), yielding an increased root mass fraction. Elevated CO₂ led to an increased relative growth rate of Larix stem basal area but no change in the final biomass of either tree species. Total understorey above‐ground mass was not altered by soil warming or elevated CO₂. However, Vaccinium myrtillus mass increased with both treatments, graminoid mass declined with warming, and forb and nonvascular plant (moss and lichen) mass decreased with both treatments. Fine roots showed a substantial reduction under soil warming (?40% for all roots <2 mm in diameter at 0–20 cm soil depth) but no change with CO₂ enrichment. Our findings suggest that enhanced overall productivity and shifts in biomass allocation will occur at the tree line, particularly with global warming. However, individual species and functional groups will respond differently to these environmental changes, with consequences for ecosystem structure and functioning.     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Changes in needle quality and larch bud moth performance in response to CO2 enrichment and defoliation of treeline larches

Abstract. 1. It is hypothesised that the larch bud moth cycle is controlled by host‐tree foliage quality. In a Free Air CO₂ Enrichment (FACE) experiment at the Swiss alpine treeline (2180 m a.s.l.), the effects of elevated CO₂ and previous year defoliation on needle quality of larch and the performance of the larch bud moth were investigated. 2. Starch and lignin concentrations increased and water content decreased in elevated CO₂‐grown needles compared with ambient CO₂ concentration. Defoliation resulted in reduced N, water, starch, and sugar concentrations in needles of the next year generation. No interactions between elevated CO₂ and defoliation on needle quality were observed. 3. Needle quality changes due to needle maturation over the course of the experiment, however, were much larger than the effects of elevated CO₂ and defoliation. For example, N concentration was on average 38% lower and lignin concentration 55% higher at the end of the experiment (early July 2003) than at the beginning (mid June 2003). 4. On non‐defoliated trees, larch bud moth larvae grew somewhat more slowly under elevated CO₂ compared with ambient CO₂ concentration. If, however, trees had been defoliated, this response was reversed, with a faster growth of larch bud moth on high CO₂‐exposed trees than on control trees. Pupal weight was not affected by either CO₂ treatment or defoliation. 5. These results suggest that the larch bud moth has to cope with large changes in food quality due to needle maturation during its development, and that additional CO₂‐ and defoliation‐induced alterations in needle chemistry have comparatively minor influences on larch bud moth performance at the treeline.     

Allocation responses to CO2 enrichment and defoliation by a native annual plant Heterotheca subaxillaris

Among plants grown under enriched atmospheric CO₂, root:shoot balance (RSB) theory predicts a proportionately greater allocation of assimilate to roots than among ambient‐grown plants. Conversely, defoliation, which decreases the plant's capacity to assimilate carbon, is predicted to increase allocation to shoot. We tested these RSB predictions, and whether responses to CO₂ enrichment were modified by defoliation, using Heterotheca subaxillaris, an annual plant native to south‐eastern USA. Plants were grown under near‐ambient (400 μmol mol^?1) and enriched (700 μmol mol^?1) levels of atmospheric CO₂. Defoliation consisted of the weekly removal of 25% of each new fully expanded, but not previously defoliated, leaf from either rosette or bolted plants. In addition to dry mass measurements of leaves, stems, and roots, Kjeldahl N, protein, starch and soluble sugars were analysed in these plant components to test the hypothesis that changes in C:N uptake ratio drive shifts in root:shoot ratio. Young, rapidly growing CO₂‐enriched plants conformed to the predictions of RSB, with higher root:shoot ratio than ambient‐grown plants (P < 0.02), whereas older, slower growing plants did not show a CO₂ effect on root:shoot ratio. Defoliation resulted in smaller plants, among which both root and shoot biomass were reduced, irrespective of CO₂ treatment (P < 0.03). However, H. subaxillaris plants were able to compensate for leaf area removal through flexible shoot allocation to more leaves vs. stem (P < 0.01). Increased carbon availability through CO₂ enrichment did not enhance the response to defoliation, apparently because of complete growth compensation for defoliation, even under ambient conditions. CO₂‐enriched plants had higher rates of photosynthesis (P < 0.0001), but this did not translate into increased final biomass accumulation. On the other hand, earlier and more abundant yield of flower biomass was an important consequence of growth under CO₂ enrichment.     

Hemiparasite abundance in an alpine treeline ecotone increases in response to atmospheric CO2 enrichment

Populations of the annual hemiparasites Melampyrum pratense L. and Melampyrum sylvaticum L. were studied at the treeline in the Swiss Alps after 3 years of in situ CO₂ enrichment. The total density of Melampyrum doubled to an average of 44 individuals per square meter at elevated CO₂ compared to ambient CO₂. In response to elevated CO₂, the height of the more abundant and more evenly distributed M. pratense increased by 20%, the number of seeds per fruit by 21%, and the total seed dry mass per fruit by 27%, but the individual seed size did not change. These results suggest that rising atmospheric CO₂ may stimulate the reproductive output and increase the abundance of Melampyrum in the alpine treeline ecotone. Because hemiparasites can have important effects on community dynamics and ecosystem processes, notably the N cycle, changing Melampyrum abundance may potentially influence the functioning of alpine ecosystems in a future CO₂-rich atmosphere.     

The carbon balance in natural and disturbed forests of the southern taiga in central Siberia

Abstract. We evaluated the balance of production and decomposition in natural ecosystems of Pinus sylvestris, Larix sibirica and Betula pendula in the southern boreal forests of central Siberia, using the Yenisei transect. We also investigated whether anthropogenic disturbances (logging, fire and recreation pressure) influence the carbon budget. Pinus and Larix stands up to age class VI act as a net sink for atmospheric carbon. Mineralization rates in young Betula forests exceed rates of uptake via photosynthesis assimilation. Old‐growth stands of all three forest types are CO₂ sources to the atmosphere. The prevalence of old‐growth Larix in the southern taiga suggests that Larix stands are a net source of CO₂. The CO₂ flux to the atmosphere exceeds the uptake of atmospheric carbon via photosynthesis by 0.23 t C.ha^‐1.yr^‐1 (47%). Betula and Pinus forests are net sinks, as photosynthesis exceeds respiration by 13% and 16% respectively. The total carbon flux from Pinus, Larix and Betula ecosystems to the atmosphere is 10 387 thousand tons C.yr^‐1. Net Primary Production (0.935 t‐C.ha^‐1) exceeds carbon release from decomposition of labile and mobile soil organic matter (Rh) by 767 thousand tons C (0.064 t‐C.ha^‐1), so that these forests are net C‐sinks. The emissions due to decomposition of slash (101 thousand tons C; 1.0%) and from fires (0.21%) are very small. The carbon balance of human‐disturbed forests is significantly different. A sharp decrease in biomass stored in Pinus and Betula ecosystems leads to decreased production. As a result, the labile organic matter pool decreased by 6–8 times; course plant residues with a low decomposition rate thus dominate this pool. Annual carbon emissions to the atmosphere from these ecosystems are determined primarily by decomposing fresh litterfall. This source comprises 40–79% of the emissions from disturbed forests compared to only 13–28% in undisturbed forests. The ratio of emissions to production (NPP) is 20–30% in disturbed and 52–76% in undisturbed forests.     

Growth response of Trifolium repens L. and Lolium perenne L. as monocultures and bi-species mixture to free air CO2 enrichment and management

Trifolium repens L. and Lolium perenne L. were grown in monocultures and bi-species mixture in a Free Air Carbon Dioxide Enrichment (FACE) experiment at elevated (60 Pa) and ambient (35 Pa) CO₂ partial pressure (pCO₂) for three years. The effects of defoliation frequencies (4 and 7 cuts in 1993; 4 and 8 cuts in 1994/95) and nitrogen fertilization (10 and 42 g m^–2 y^–1 N in 1993; 14 and 56 g m^–2 y^–1 N in 1994/95) on the growth response to pCO₂ were investigated. There were significant interspecific differences in the CO₂ responses during the first two years, while in the third growing season, these interspecific differences disappeared. Yield of T. repens in monocultures increased in the first two years by 20% when grown at elevated pCO₂. This CO₂ response was independent of defoliation frequency and nitrogen fertilization. In the third year, the CO₂ response of T. repens declined to 11%. In contrast, yield of L. perenne monocultures increased by only 7% on average over three years at elevated pCO₂. The yield response of L. perenne to CO₂ changed according to defoliation frequency and nitrogen fertilization, mainly in the second and third year. The ratio of root/yield of L. perenne increased under elevated pCO₂, low N fertilizer rate, and frequent defoliation, but it remained unchanged in T. repens. We suggest that the more abundant root growth of L. perenne was related to increased N limitation under elevated pCO₂. The consequence of these interspecific differences in the CO₂ response was a higher proportion of T. repens in the mixed swards at elevated pCO₂. This was evident in all combinations of defoliation and nitrogen treatments. However, the proportion of the species was more strongly affected by N fertilization and defoliation frequency than by elevated pCO₂. Based on these results, we conclude that the species proportion in managed grassland may change as the CO₂ concentration increases. However, an adapted management could, at least partially, counteract such CO₂ induced changes in the proportion of the species. Since the availability of mineral N in the soil may be important for the species’ responses to elevated pCO₂, more long-term studies, particularly of processes in the soil, are required to predict the entire ecosystem response.     

Effect of elevated [CO2] on stem wood properties of mature Norway spruce grown at different soil nutrient availability

The objective of the present study was to investigate the interactive effects of elevated [CO₂] and soil nutrient availability on secondary xylem structure and chemical composition of 41‐year‐old Norway spruce (Picea abies (L.) Karst.) trees. The nonfertilized and irrigated‐fertilized trees were, for 3 years, continuously exposed to elevated [CO₂] in whole‐tree chambers. Elevated [CO₂] decreased concentrations of soluble sugars, acid‐soluble lignin and nitrogen in stem wood, but the effects were not consistent between sampling height and/or fertilization. The effect of 2*ambient [CO₂] on wood structure depended on the exposure year and/or fertilization. Radial lumen diameter decreased and annual ring width increased in the second year of exposure (1999) in elevated [CO₂]. In the latter, the CO₂ effect was significant only in the nonfertilized trees. Stem wood chemistry and structure were significantly affected by fertilization. Fertilization increased the concentrations of nitrogen and gravimetric lignin, annual ring width, and radial lumen diameter. Fertilization decreased C/N ratio, mean ring density, earlywood density, latewood density, cell wall thickness, cell wall index, and latewood percentage. We conclude that elevated [CO₂] had only minor effects on wood properties while fertilization had more marked effects and thus may affect ecosystem processes and suitability of wood for different end‐use purposes.     

Interannual climatic variation mediates elevated CO2 and O3 effects on forest growth

We analyzed growth data from model aspen (Populus tremuloides Michx.) forest ecosystems grown in elevated atmospheric carbon dioxide ([CO₂]; 518 μL L^?1) and ozone concentrations ([O₃]; 1.5 × background of 30–40 nL L^?1 during daylight hours) for 7 years using free‐air CO₂ enrichment technology to determine how interannual variability in present‐day climate might affect growth responses to either gas. We also tested whether growth effects of those gasses were sustained over time. Elevated [CO₂] increased tree heights, diameters, and main stem volumes by 11%, 16%, and 20%, respectively, whereas elevated ozone [O₃] decreased them by 11%, 8%, and 29%, respectively. Responses similar to these were found for stand volume and basal area. There were no growth responses to the combination of elevated [CO₂+O₃]. The elevated [CO₂] growth stimulation was found to be decreasing, but relative growth rates varied considerably from year to year. Neither the variation in annual relative growth rates nor the apparent decline in CO₂ growth response could be explained in terms of nitrogen or water limitations. Instead, growth responses to elevated [CO₂] and [O₃] interacted strongly with present‐day interannual variability in climatic conditions. The amount of photosynthetically active radiation and temperature during specific times of the year coinciding with growth phenology explained 20–63% of the annual variation in growth response to elevated [CO₂] and [O₃]. Years with higher photosynthetic photon flux (PPF) during the month of July resulted in more positive growth responses to elevated [CO₂] and more negative growth responses to elevated [O₃]. Mean daily temperatures during the month of October affected growth in a similar fashion the following year. These results indicate that a several‐year trend of increasingly cloudy summers and cool autumns were responsible for the decrease in CO₂ growth response.     

Non-structural carbon compounds in temperate forest trees

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of non‐structural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad‐leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring‐porous versus diffuse‐porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris. The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues (‘fat trees’; Tilia, Pinus, Picea, Larix), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO₂ enrichment. The classical view that deciduous trees rely more on C‐reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO₂ concentrations over the last 150 years.     

Effects of atmospheric CO2 enrichment on early growth of Vivia faba, a plant with large cotyledons

Seedlings of Vicia faba L. were grown in open-top growth chambers at present (P=350μmol^?1) and at elevated (E=700μmol mol^?1) atmospheric CO₂ concentration. The effects of CO₂ enrichment on the first phase of growth after germination were examined over 45 d. There were no positive effects of CO₂ enrichment on growth of the seedlings during this early phase. No differences were observed in leaf area or in total dry weight. No differences were found in morphology or anatomy of the leaves. The numbers of stomatal and epidermal cells, thickness of leaf, of epidermis and of mesophyll cell-layers were unaffected by CO₂ enrichment. Also, no differences were observed in leaf concentrations of chlorophyll, reducing carbohydrates or starch. These results contrast markedly with results from similar experiments on poplar hybrids and Phaseolus vulgaris obtained in the same growth facility. It seems that the intitial growth is under internal control such that the atmospheric CO₂ concentration has no effects. The lack of response in this case may be attributed to the presence and longevity of the large cotyledons which provided available substrate for growth.     

Growth and phenology of mature temperate forest trees in elevated CO2

Are mature forest trees carbon limited at current CO₂ concentrations? Will ‘mid‐life’, 35 m tall deciduous trees grow faster in a CO₂‐enriched atmosphere? To answer these questions we exposed ca. 100‐year‐old temperate forest trees at the Swiss Canopy Crane site near Basel, Switzerland to a ca. 540 ppm CO₂ atmosphere using web‐FACE technology. Here, we report growth responses to elevated CO₂ for 11 tall trees (compared with 32 controls) of five species during the initial four treatment years. Tested across all trees, there was no CO₂ effect on stem basal area (BA) increment (neither when tested per year nor cumulatively for 4 years). In fact, the 4th year means were almost identical for the two groups. Stem growth data were standardized by pretreatment growth (5 years) in order to account for a priori individual differences in vigor. Although this experiment was not designed to test species specific effects, one species, the common European beech, Fagus sylvatica, showed a significant growth enhancement in the first year, which reoccurred during a centennial drought in the third year. None of the other dominant species (Quercus petraea, Carpinus betulus) showed a growth response to CO₂ in any of the 4 years or for all years together. The inclusion or exclusion of single individuals of Prunus avium and Tilia platyphyllos did not change the picture. In elevated CO₂, lateral branching in terminal shoots was higher in Fagus in 2002, when shoots developed from buds that were formed during the first season of CO₂ enrichment (2001), but there was no effect in later years and no change in lateral branching in any of the other species. In Quercus, there was a steady stimulation of leading shoot length in high‐CO₂ trees. Phenological variables (bud break, leaf fall, leaf duration) were highly species specific and were not affected by elevated CO₂ in any consistent way. Our 4‐year data set reflects a very dynamic and species‐specific response of tree growth to a step change in CO₂ supply. Stem growth after 4 years of exposure does not support the notion that mature forest trees will accrete wood biomass at faster rates in a future CO₂‐enriched atmosphere.     

Defoliation reduces soil biota – and modifies stimulating effects of elevated CO2

To understand the responses to external disturbance such as defoliation and possible feedback mechanisms at global change in terrestrial ecosystems, it is necessary to examine the extent and nature of effects on aboveground–belowground interactions. We studied a temperate heathland system subjected to experimental climate and atmospheric factors based on prognoses for year 2075 and further exposed to defoliation. By defoliating plants, we were able to study how global change modifies the interactions of the plant–soil system. Shoot production, root biomass, microbial biomass, and nematode abundance were assessed in the rhizosphere of manually defoliated patches of Deschampsia flexuosa in June in a full‐factorial FACE experiment with the treatments: increased atmospheric CO₂, increased nighttime temperatures, summer droughts, and all of their combinations. We found a negative effect of defoliation on microbial biomass that was not apparently affected by global change. The negative effect of defoliation cascades through to soil nematodes as dependent on CO₂ and drought. At ambient CO₂, drought and defoliation each reduced nematodes. In contrast, at elevated CO₂, a combination of drought and defoliation was needed to reduce nematodes. We found positive effects of CO₂ on root density and microbial biomass. Defoliation affected soil biota negatively, whereas elevated CO₂ stimulated the plant–soil system. This effect seen in June is contrasted by the effects seen in September at the same site. Late season defoliation increased activity and biomass of soil biota and more so at elevated CO₂. Based on soil biota responses, plants defoliated in active growth therefore conserve resources, whereas defoliation after termination of growth results in release of resources. This result challenges the idea that plants via exudation of organic carbon stimulate their rhizosphere biota when in apparent need of nutrients for growth.     

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO₂ levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO₂ enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO₂ and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO₂ levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.     

Nitrogen-regulated effects of free-air CO2 enrichment on methane emissions from paddy rice fields

Using the free‐air CO₂ enrichment (FACE) techniques, we carried out a 3‐year mono‐factorial experiment in temperate paddy rice fields of Japan (1998–2000) and a 3‐year multifactorial experiment in subtropical paddy rice fields in the Yangtze River delta in China (2001–2003), to investigate the methane (CH₄) emissions in response to an elevated atmospheric CO₂ concentration (200±40 mmol mol^?1 higher than that in the ambient atmosphere). No significant effect of the elevated CO₂ upon seasonal accumulative CH₄ emissions was observed in the first rice season, but significant stimulatory effects (CH₄ increase ranging from 38% to 188%, with a mean of 88%) were observed in the second and third rice seasons in the fields with or without organic matter addition. The stimulatory effects of the elevated CO₂ upon seasonal accumulative CH₄ emissions were negatively correlated with the addition rates of decomposable organic carbon (P<0.05), but positively with the rates of nitrogen fertilizers applied in either the current rice season (P<0.05) or the whole year (P<0.01). Six mechanisms were proposed to explain collectively the observations. Soil nitrogen availability was identified as an important regulator. The effect of soil nitrogen availability on the observed relation between elevated CO₂ and CH₄ emission can be explained by (a) modifying the C/N ratio of the plant residues formed in the previous growing season(s); (b) changing the inhibitory effect of high C/N ratio on plant residue decomposition in the current growing season; and (c) altering the stimulatory effects of CO₂ enrichment upon plant growth, as well as nitrogen uptake in the current growing season. This study implies that the concurrent enrichment of reactive nitrogen in the global ecosystems may accelerate the increase of atmospheric methane by initiating a stimulatory effect of the ongoing dramatic atmospheric CO₂ enrichment upon methane emissions from nitrogen‐poor paddy rice ecosystems and further amplifying the existing stimulatory effect in nitrogen‐rich paddy rice ecosystems.     

The microfood web of grassland soils responds to a moderate increase in atmospheric CO2

The response of the soil microfood web (microflora, nematodes) to a moderate increase in atmospheric CO₂ (+20%) was investigated by means of a free air CO₂ enrichment experiment. The study was carried out in a seminatural temperate grassland for a period of 4 consecutive years (1 year before fumigation commenced and 3 years with fumigation). Several soil biological parameters showed no change (microbial biomass, bacterial biomass) or decline (microbial respiration) in the first year of elevated CO₂ treatment as compared with controls. Each of these parameters were higher than controls, however, after 3 years of treatment. The relative abundance of predaceous nematodes also decreased in year 1 of the experiment, increased in year 2, but decreased again in year 3. In contrast, the relative abundance of root hair feeding nematodes, at first, increased under elevated CO₂ and then returned to the initial level again. Increased microbial biomass indicates enhanced C storage in the labile carbon pool of the active microfood web in subsequent years. According to measurements on the amounts of soil extractable C, changes in resource availability seem to be key to the response of the soil microfood web. We found a strong response of bacteria to elevated CO₂, while the fungal biomass remained largely unchanged. This contrasts to findings reported in the literature. We hypothesize that this may be because of contrasting effects of different levels of CO₂ enrichment on the microbial community (i.e. stimulation of bacteria at moderate levels and stimulation of fungi at high levels of CO₂ enrichment). However, various CO₂ effects observed in our study are similar in magnitude to those observed in other studies for a much higher level of atmospheric carbon. These include the particular sensitivity of predaceous nematodes and the long‐term increase of microbial respiration. Our findings confirm that the potential of terrestrial ecosystems to accumulate additional carbon might be lower than previously thought. Furthermore, CO₂‐induced changes of temperate grassland ecosystems might emerge much earlier than expected.     

Fine root dynamics in a loblolly pine forest are influenced by free-air-CO2-enrichment: a six-year-minirhizotron study

Efforts to characterize carbon (C) cycling among atmosphere, forest canopy, and soil C pools are hindered by poorly quantified fine root dynamics. We characterized the influence of free‐air‐CO₂‐enrichment (ambient +200 ppm) on fine roots for a period of 6 years (Autumn 1998 through Autumn 2004) in an 18‐year‐old loblolly pine (Pinus taeda) plantation near Durham, NC, USA using minirhizotrons. Root production and mortality were synchronous processes that peaked most years during spring and early summer. Seasonality of fine root production and mortality was not influenced by atmospheric CO₂ availability. Averaged over all 6 years of the study, CO₂ enrichment increased average fine root standing crop (+23%), annual root length production (+25%), and annual root length mortality (+36%). Larger increase in mortality compared with production with CO₂ enrichment is explained by shorter average fine root lifespans in elevated plots (500 days) compared with controls (574 days). The effects of CO₂‐enrichment on fine root proliferation tended to shift from shallow (0–15 cm) to deeper soil depths (15–30) with increasing duration of the study. Diameters of fine roots were initially increased by CO₂‐enrichment but this effect diminished over time. Averaged over 6 years, annual fine root NPP was estimated to be 163 g dw m^?2 yr^?1 in CO₂‐enriched plots and 130 g dw m^?2 yr^?1 in control plots (P= 0.13) corresponding to an average annual additional input of fine root biomass to soil of 33 g m^?2 yr^?1 in CO₂‐enriched plots. A lack of consistent CO₂× year effects suggest that the positive effects of CO₂ enrichment on fine root growth persisted 6 years following minirhizotron tube installation (8 years following initiation of the CO₂ fumigation). Although CO₂‐enrichment contributed to extra flow of C into soil in this experiment, the magnitude of the effect was small suggesting only modest potential for fine root processes to directly contribute to soil C storage in south‐eastern pine forests.     

Growth and reproduction of the alpine grasshopper <Emphasis Type="Italic">Miramella alpina</Emphasis> feeding on CO<Subscript>2</Subscript>-enriched dwarf shrubs at treeline

The consequences for plant-insect interactions of atmospheric changes in alpine ecosystems are not well understood. Here, we tested the effects of elevated CO₂ on leaf quality in two dwarf shrub species (Vaccinium myrtillus and V. uliginosum) and the response of the alpine grasshopper (Miramella alpina) feeding on these plants in a field experiment at the alpine treeline (2,180 m a.s.l.) in Davos, Switzerland. Relative growth rates (RGR) of M. alpina nymphs were lower when they were feeding on V. myrtillus compared to V. uliginosum, and were affected by elevated CO₂ depending on plant species and nymph developmental stage. Changes in RGR correlated with CO₂-induced changes in leaf water, nitrogen, and starch concentrations. Elevated CO₂ resulted in reduced female adult weight irrespective of plant species, and prolonged development time on V. uliginosum only, but there were no significant differences in nymphal mortality. Newly molted adults of M. alpina produced lighter eggs and less secretion (serving as egg protection) under elevated CO₂. When grasshoppers had a choice among four different plant species grown either under ambient or elevated CO₂, V. myrtillus and V. uliginosum consumption increased under elevated CO₂ in females while it decreased in males compared to ambient CO₂-grown leaves. Our findings suggest that rising atmospheric CO₂ distinctly affects leaf chemistry in two important dwarf shrub species at the alpine treeline, leading to changes in feeding behavior, growth, and reproduction of the most important insect herbivore in this system. Changes in plant-grasshopper interactions might have significant long-term impacts on herbivore pressure, community dynamics and ecosystem stability in the alpine treeline ecotone.