Fitness of natural willow hybrids in a pioneer mosaic hybrid zone

Hybrid fitness is an important parameter to predict the evolutionary consequences of a hybridization event and to characterize hybrid zones. We studied fitness parameters of F₁ and later‐generation hybrids between the lowland species Salix purpurea and the alpine S. helvetica that have recently emerged during colonization of an alpine glacier forefield. Fruit production (number of capsules per catkin and fruit set) did not differ between hybrids and parents, but the number of seeds per capsule of F₁ hybrids was slightly lower than that of later‐generation hybrids and of the parents. Germination rates and seedling growth were tested on three substrates (pH 4.5, 7.0, and 8.0). Germination rates of seeds collected from F₁ hybrids were lower on acid and neutral substrates, but equal at pH 8.0 compared to all other groups, while the seeds from later‐generation hybrids performed as well as the parents on all three substrates. In seedling growth, the colonizer S. purpurea performed better than all other taxa on all three substrates, while hybrids resembled the subalpine species S. helvetica. Results suggest that endogenous selection acts against F₁ hybrids, but favors fitter genotypes in later‐generation hybrids. Exogenous selection via soil pH appears to be weak during seedling establishment. The pioneer vegetation on the glacier forefield may offer sufficient niche space for hybrid seedlings. Owing to the relatively high fitness of the hybrids and the scattered distribution of hybrids and parental individuals on the glacier forefield, this hybrid zone can be assigned to a mosaic model, probably facilitating gene flow and introgression between the parental species. As establishment of the hybrid zone appears to be linked to a colonization process, we propose to call it a pioneer mosaic hybrid zone.     

Species-specific responses of plant communities to altered carbon and nutrient availability

In a field microcosm experiment, species‐specific responses of aboveground biomass of two California annual grassland communities to elevated CO₂ and nutrient availability were investigated. One community grows on shallow, nutrient‐poor serpentine‐derived soil whereas the other occurs on deeper, modestly fertile sandstone/greenstone‐derived substrate. In most species, CO₂ effects did not appear until late in the growing season, probably because the elevated CO₂ increased water‐use‐efficiency easing, the onset of the summer drought. Responses of aboveground biomass to elevated CO₂ differed depending on nutrient availability. Similarly, biomass responses to nutrient treatments differed depending on the CO₂ status. For the majority of the species, production increased most under elevated CO₂ with added nutrients (N,P,K, and micro nutrients). Some species were losers under conditions that increased overall community production, including Bromus hordeaceus in the serpentine community (negative biomass response under elevated CO₂) and Lotus wrangelianus in both communities (negative biomass response with added nitrogen). Treatment and competitive effects on species‐specific biomass varied in both magnitude and direction, especially in the serpentine community, significantly affecting community structure. Individual resource environments are likely to be affected by neighbouring plants, and these competitive interactions complicate predictions of species' responses to elevated CO₂.     

Elevated atmospheric CO2 stimulates aboveground biomass in a fire-regenerated scrub-oak ecosystem

The effect of elevated atmospheric CO₂ concentration (C_a) on the aboveground biomass of three oak species, Quercus myrtifolia, Q. geminata, and Q. chapmanii, was estimated nondestructively using allometric relationships between stem diameter and aboveground biomass after four years of experimental treatment in a naturally fire‐regenerated scrub‐oak ecosystem. After burning a stand of scrub‐oak vegetation, re‐growing plants were exposed to either current ambient (379 µL L^?1 CO₂) or elevated (704 µL L^?1 CO₂) C_a in 16 open‐top chambers over a four‐year period, and measurements of stem diameter were carried out annually on all oak shoots within each chamber. Elevated C_a significantly increased aboveground biomass, expressed either per unit ground area or per shoot; elevated C_a had no effect on shoot density. The relative effect of elevated C_a on aboveground biomass increased each year of the study from 44% (May 96–Jan 97), to 55% (Jan 97–Jan 98), 66% (Jan 98–Jan 99), and 75% (Jan 99–Jan 00). The effect of elevated C_a was species specific: elevated C_a significantly increased aboveground biomass of the dominant species, Q. myrtifolia, and tended to increase aboveground biomass of Q. chapmanii, but had no effect on aboveground biomass of the subdominant, Q. geminata. These results show that rising atmospheric CO₂ has the potential to stimulate aboveground biomass production in ecosystems dominated by woody species, and that species‐specific growth responses could, in the long term, alter the composition of the scrub‐oak community.     

Tree species diversity interacts with elevated CO2 to induce a greater root system response

As a consequence of land‐use change and the burning of fossil fuels, atmospheric concentrations of CO₂ are increasing and altering the dynamics of the carbon cycle in forest ecosystems. In a number of studies using single tree species, fine root biomass has been shown to be strongly increased by elevated CO₂. However, natural forests are often intimate mixtures of a number of co‐occurring species. To investigate the interaction between tree mixture and elevated CO₂, Alnus glutinosa, Betula pendula and Fagus sylvatica were planted in areas of single species and a three species polyculture in a free‐air CO₂ enrichment study (BangorFACE). The trees were exposed to ambient or elevated CO₂ (580 μmol mol^?1) for 4 years. Fine and coarse root biomass, together with fine root turnover and fine root morphological characteristics were measured. Fine root biomass and morphology responded differentially to the elevated CO₂ at different soil depths in the three species when grown in monocultures. In polyculture, a greater response to elevated CO₂ was observed in coarse roots to a depth of 20 cm, and fine root area index to a depth of 30 cm. Total fine root biomass was positively affected by elevated CO₂ at the end of the experiment, but not by species diversity. Our data suggest that existing biogeochemical cycling models parameterized with data from species grown in monoculture may be underestimating the belowground response to global change.     

Partitioning of ecosystem respiration of CO2 released during land‐use transition from temperate agricultural grassland to Miscanthus × giganteus

Conversion of large areas of agricultural grassland is inevitable if European and UK domestic production of biomass is to play a significant role in meeting demand. Understanding the impact of these land‐use changes on soil carbon cycling and stocks depends on accurate predictions from well‐parameterized models. Key considerations are cultivation disturbance and the effect of autotrophic root input stimulation on soil carbon decomposition under novel biomass crops. This study presents partitioned parameters from the conversion of semi‐improved grassland to Miscanthus bioenergy production and compares the contribution of autotrophic and heterotrophic respiration to overall ecosystem respiration of CO₂ in the first and second years of establishment. Repeated measures of respiration from within and without root exclusion collars were used to produce time‐series model integrations separating live root inputs from decomposition of grass residues ploughed in with cultivation of the new crop. These parameters were then compared to total ecosystem respiration derived from eddy covariance sensors. Average soil surface respiration was 13.4% higher in the second growing season, increasing from 2.9 to 3.29 g CO₂‐C m^?2 day^?1. Total ecosystem respiration followed a similar trend, increasing from 4.07 to 5.4 g CO₂‐C m^?2 day^?1. Heterotrophic respiration from the root exclusion collars was 32.2% lower in the second growing season at 1.20 g CO₂‐C m^?2 day^?1 compared to the previous year at 1.77 g CO₂‐C m^?2 day^?1. Of the total respiration flux over the two‐year time period, aboveground autotrophic respiration plus litter decomposition contributed 38.46% to total ecosystem respiration while belowground autotrophic respiration and stimulation by live root inputs contributed 46.44% to soil surface respiration. This figure is notably higher than mean figures for nonforest soils derived from the literature and demonstrates the importance of crop‐specific parameterization of respiration models.     

Comparison of the effect of elevated CO2 on an invasive species (Centaurea solstitialis) in monoculture and community settings

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) is likely to change the species composition of plant communities. To investigate whether growth of a highly invasive plant species, Centaurea solstitialis (yellow starthistle), was affected by elevated [CO₂], and whether the success of this species would increase under CO₂ enrichment, I grew the species in serpentine soil microcosms, both as a monoculture and as a component of a grassland community. Centaurea grown in monoculture responded strongly to [CO₂] enrichment of 350 mol mol^–1, increasing aboveground biomass production by 70%, inflorescence production by 74%, and midday photosynthesis by an average of 132%. When grown in competition with common serpentine grassland species, Centaurea responded to CO₂ enrichment with similar but nonsignificant increases (+69% aboveground biomass, +71% inflorescence production), while total aboveground biomass of the polyculture increased by 28%. Centaurea's positive CO₂ response in monoculture and parallel (but non-significant) response in polyculture provoke questions about possible consequences of increasing CO₂ for more typical California grasslands, where the invader already causes major problems.     

Climate‐sensitive ecosystem carbon dynamics along the soil chronosequence of the Damma glacier forefield,Switzerland

We performed a detailed study on the carbon build‐up over the 140‐year‐long chronosequence of the Damma glacier forefield, Switzerland, to gain insights into the organic carbon dynamics during the initial stage of soil formation and ecosystem development. We determined soil carbon and nitrogen contents and their stable isotopic compositions, as well as molecular‐level composition of the bulk soils, and recalcitrance parameters of carbon in different fractions. The chronosequence was divided into three age groups, separated by small end moraines that resulted from two glacier re‐advances. The net ecosystem carbon balance (NECB) showed an exponential increase over the last decades, with mean annual values that range from 100 g C m^?2 yr^?1 in the youngest part to over 300 g C m^?2 yr^?1 in a 60–80 years old part. However, over the entire 140‐year chronosequence, the NECB is only 20 g C m^?2 yr^?1, similar to results of other glacier forefield studies. The difference between the short‐ and long‐term NECB appears to be caused by reductions in ecosystem carbon (EC) accumulation during periods with a colder climate. We propose that two complementary mechanisms have been responsible: 1) Reductions in net primary productivity down to 50% below the long‐term mean, which we estimated using reconstructed effective temperature sums. 2) Disturbance of sites near the terminus of the re‐advanced glacier front. Stabilization of soil organic matter appeared to play only a minor role in the coarse‐grained forefield. We conclude that the forefield ecosystem, especially primary productivity, reacts rapidly to climate changes. The EC gained at warm periods is easily lost again in a cooling climate. Our conclusions may also be valid for other high mountain ecosystems and possibly arctic ecosystems.     

Invasive submerged freshwater macrophytes are more plastic in their response to light intensity than to the availability of free CO2 in air‐equilibrated water

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Offsetting high water demands with high productivity: Sorghum as a biofuel crop in a high irradiance arid ecosystem

High irradiance arid environments are promising, yet understudied, areas for biofuel production. We investigated the productivity and environmental trade‐offs of growing sorghum (Sorghum bicolor) as a biofuel feedstock in the low deserts of California (CA). Using a 5.3 ha experimental field in the Imperial Valley, CA, we measured aboveground biomass production and net ecosystem exchange of CO₂ and H₂O via eddy covariance over three growing periods between February and November 2012. Environmental conditions were extreme, with high irradiance, vapor pressure deficit (VPD), and air temperature throughout the growing season. Air temperature peaked in August with a maximum of 45.7 °C. Sorghum produced an annual aboveground biomass yield of 43.7 Mg per hectare. Net ecosystem exchange (NEE) was highest during the summer growth period and reached a maximum of ?68 μmol CO₂ m^?2 s^?1. Water use efficiency, or biomass water ratio (BWR), was high (4.0 g dry biomass kg^?1 H₂O) despite high seasonal evapotranspiration (1094 kg H₂O m^?2). The BWR of sorghum surpassed that of many C4 biofuel candidate crops in the United States, as well as that of alfalfa which is currently widely grown in the Imperial Valley. Sorghum also outperformed many US biofuel crops in terms of radiation use efficiency (RUE), achieving 1.5 g dry biomass MJ^?1. We found no evidence of saturation of NEE at high levels of photosynthetically active radiation (PAR) (up to 2250 μmol m^?2 s^?1). In addition, we found no evidence that NEE was inhibited by either high VPD or air temperature during peak photosynthetic phases. The combination of high productivity, high BWR, and high RUE suggests that sorghum is well adapted to this extreme environment. The biomass production rates and efficiency metrics spanning three growing periods provide fundamental data for future Life Cycle Assessments (LCA), which are needed to assess the sustainability of this sorghum biofuel feedstock system.     

Interactive effects of nitrate concentrations and carbon dioxide on the stoichiometry,biomass allocation and growth rate of submerged aquatic plants

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Temperature and precipitation drive temporal variability in aquatic carbon and GHG concentrations and fluxes in a peatland catchment

The aquatic pathway is increasingly being recognized as an important component of catchment carbon and greenhouse gas (GHG) budgets, particularly in peatland systems due to their large carbon store and strong hydrological connectivity. In this study, we present a complete 5‐year data set of all aquatic carbon and GHG species from an ombrotrophic Scottish peatland. Measured species include particulate and dissolved forms of organic carbon (POC, DOC), dissolved inorganic carbon (DIC), CO₂, CH₄ and N₂O. We show that short‐term variability in concentrations exists across all species and this is strongly linked to discharge. Seasonal cyclicity was only evident in DOC, CO₂ and CH₄ concentration; however, temperature correlated with monthly means in all species except DIC. Although the temperature correlation with monthly DOC and POC concentrations appeared to be related to biological productivity in the terrestrial system, we suggest the temperature correlation with CO₂ and CH₄ was primarily due to in‐stream temperature‐dependent solubility. Interannual variability in total aquatic carbon concentration was strongly correlated with catchment gross primary productivity (GPP) indicating a strong potential terrestrial aquatic linkage. DOC represented the largest aquatic carbon flux term (19.3 ± 4.59 g C m^?2 yr^?1), followed by CO₂ evasion (10.0 g C m^?2 yr^?1). Despite an estimated contribution to the total aquatic carbon flux of between 8 and 48%, evasion estimates had the greatest uncertainty. Interannual variability in total aquatic carbon export was low in comparison with variability in terrestrial biosphere–atmosphere exchange, and could be explained primarily by temperature and precipitation. Our results therefore suggest that climatic change is likely to have a significant impact on annual carbon losses through the aquatic pathway, and as such, aquatic exports are fundamental to the understanding of whole catchment responses to climate change.     

Canopy warming caused photosynthetic acclimation and reduced seed yield in maize grown at ambient and elevated [CO2]

Rising atmospheric CO₂ concentration ([CO₂]) and attendant increases in growing season temperature are expected to be the most important global change factors impacting production agriculture. Although maize is the most highly produced crop worldwide, few studies have evaluated the interactive effects of elevated [CO₂] and temperature on its photosynthetic physiology, agronomic traits or biomass, and seed yield under open field conditions. This study investigates the effects of rising [CO₂] and warmer temperature, independently and in combination, on maize grown in the field throughout a full growing season. Free‐air CO₂ enrichment (FACE) technology was used to target atmospheric [CO₂] to 200 μmol mol^?1 above ambient [CO₂] and infrared heaters to target a plant canopy increase of 3.5 °C, with actual season mean heating of ~2.7 °C, mimicking conditions predicted by the second half of this century. Photosynthetic gas‐exchange parameters, leaf nitrogen and carbon content, leaf water potential components, and developmental measurements were collected throughout the season, and biomass and yield were measured at the end of the growing season. As predicted for a C₄ plant, elevated [CO₂] did not stimulate photosynthesis, biomass, or yield. Canopy warming caused a large shift in aboveground allocation by stimulating season‐long vegetative biomass and decreasing reproductive biomass accumulation at both CO₂ concentrations, resulting in decreased harvest index. Warming caused a reduction in photosynthesis due to down‐regulation of photosynthetic biochemical parameters and the decrease in the electron transport rate. The reduction in seed yield with warming was driven by reduced photosynthetic capacity and by a shift in aboveground carbon allocation away from reproduction. This field study portends that future warming will reduce yield in maize, and this will not be mitigated by higher atmospheric [CO₂] unless appropriate adaptation traits can be introduced into future cultivars.     

Flowering in grassland predicted by CO2 and resource effects on species aboveground biomass

Continuing enrichment of atmospheric CO₂ may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO₂ enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C₄‐dominated grassland communities on contrasting soils along a CO₂ concentration gradient spanning pre‐industrial to expected mid‐21st century levels (250–500 μl/L). CO₂ enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO₂ enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R² = .34–.74), a measure of species abundance. CO₂ enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species‐specific controls on flowering in two general forms: (1) CO₂ effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO₂ effect. Understanding the heterogeneity in species responses to CO₂ enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO₂ effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes.     

Simple additive effects are rare: a quantitative review of plant biomass and soil process responses to combined manipulations of CO2 and temperature

In recent years, increased awareness of the potential interactions between rising atmospheric CO₂ concentrations ([ CO₂ ]) and temperature has illustrated the importance of multifactorial ecosystem manipulation experiments for validating Earth System models. To address the urgent need for increased understanding of responses in multifactorial experiments, this article synthesizes how ecosystem productivity and soil processes respond to combined warming and [ CO₂ ] manipulation, and compares it with those obtained in single factor [ CO₂ ] and temperature manipulation experiments. Across all combined elevated [ CO₂ ] and warming experiments, biomass production and soil respiration were typically enhanced. Responses to the combined treatment were more similar to those in the [ CO₂ ]‐only treatment than to those in the warming‐only treatment. In contrast to warming‐only experiments, both the combined and the [ CO₂ ]‐only treatments elicited larger stimulation of fine root biomass than of aboveground biomass, consistently stimulated soil respiration, and decreased foliar nitrogen (N) concentration. Nonetheless, mineral N availability declined less in the combined treatment than in the [ CO₂ ]‐only treatment, possibly due to the warming‐induced acceleration of decomposition, implying that progressive nitrogen limitation (PNL) may not occur as commonly as anticipated from single factor [ CO₂ ] treatment studies. Responses of total plant biomass, especially of aboveground biomass, revealed antagonistic interactions between elevated [ CO₂ ] and warming, i.e. the response to the combined treatment was usually less‐than‐additive. This implies that productivity projections might be overestimated when models are parameterized based on single factor responses. Our results highlight the need for more (and especially more long‐term) multifactor manipulation experiments. Because single factor CO₂ responses often dominated over warming responses in the combined treatments, our results also suggest that projected responses to future global warming in Earth System models should not be parameterized using single factor warming experiments.     

Agricultural greenhouse gas mitigation potential globally,in Europe and in the UK: what have we learnt in the last 20 years?

Agricultural lands occupy about 40–50% of the Earth's land surface. Agricultural practices can make a significant contribution at low cost to increasing soil carbon sinks, reducing greenhouse gas (GHG) emissions and contributing biomass feedstocks for energy use. Considering all gases, the global technical mitigation potential from agriculture (excluding fossil fuel offsets from biomass) by 2030 is estimated to be ca. 5500–6000 Mt CO₂‐eq. yr^?1. Economic potentials are estimated to be 1500–1600, 2500–2700 and 4000–4300 Mt CO₂‐eq. yr^?1 at carbon prices of up to $US20, 50 and 100 t CO₂‐eq.^?1, respectively. The value of the global agricultural GHG mitigation at the same three carbon prices is $US32 000, 130 000 and 420 000 million yr^?1, respectively. At the European level, early estimates of soil carbon sequestration potential in croplands were ca. 200 Mt CO₂ yr^?1, but this is a technical potential and is for geographical Europe as far east as the Urals. The economic potential is much smaller, with more recent estimates for the EU27 suggesting a maximum potential of ca. 20 Mt CO₂‐eq. yr^?1. The UK is small in global terms, but a large part of its land area (11 Mha) is used for agriculture. Agriculture accounts for about 7% of total UK GHG emissions. The mitigation potential of UK agriculture is estimated to be ca. 1–2 Mt CO₂‐eq. yr^?1, accounting for less than 1% of UK total GHG emissions.     

Transpiration of a 64-year old maritime pine stand in Portugal

Alpine plant species have been shown to exhibit a more pronounced increase in leaf photosynthesis under elevated CO₂ than lowland plants. In order to test whether this higher carbon fixation efficiency will translate into increased biomass production under CO₂ enrichment we exposed plots of narrow alpine grassland (Swiss Central Alps, 2470 m) to ambient (355 μl l^-1) and elevated (680 μl l^-1) CO₂ concentration using open top chambers. Part of the plost received moderate mineral nutrient additions (40 kg ha^-1 year^-1 of nitrogen in a complete fertilizer mix). Under natural nutrient supply CO₂ enrichment had no effect on biomass production per unit land area during any of the three seasons studied so far. Correspondingly, the dominant species Carex curvula and Leontodon helveticus as well as Trifolium alpinum did not show a growth response either at the population level or at the shoot level. However, the subdominant generalistic species Poa alpina strongly increased shoot growth (+47%). Annual root production (in ingrowth cores) was significantly enhanced in C. curvula in the 2nd and 3rd year of investigation (+43%) but was not altered in the bulk samples for all species. Fertilizer addition generally stimulated above-ground (+48%) and below-ground (+26%) biomass production right from the beginning. Annual variations in weather conditions during summer also strongly influenced above-ground biomass production (19–27% more biomass in warm seasons compared to cool seasons). However, neither nutrient availability nor climate had a significant effect on the CO₂ response of the plants. Our results do not support the hypothesis that alpine plants, due to their higher carbon uptake efficiency, will increase biomass production under future atmospheric CO₂ enrichment, at least not in such late successional communities. However, as indicated by the response of P. alpina, species-specific responses occur which may lead to altered community structure and perhaps ecosystem functioning in the long-term. Our findings further suggest that possible climatic changes are likely to have a greater impact on plant growth in alpine environments than the direct stimulation of photosynthesis by CO₂. Counter-intuitively, our results suggest that even under moderate climate warming or enhanced atmospheric nitrogen deposition positive biomass responses to CO₂ enrichment of the currently dominating species are unlikely.     

Impacts of 3 years of elevated atmospheric CO2 on rhizosphere carbon flow and microbial community dynamics

Carbon (C) uptake by terrestrial ecosystems represents an important option for partially mitigating anthropogenic CO₂ emissions. Short‐term atmospheric elevated CO₂ exposure has been shown to create major shifts in C flow routes and diversity of the active soil‐borne microbial community. Long‐term increases in CO₂ have been hypothesized to have subtle effects due to the potential adaptation of soil microorganism to the increased flow of organic C. Here, we studied the effects of prolonged elevated atmospheric CO₂ exposure on microbial C flow and microbial communities in the rhizosphere. Carex arenaria (a nonmycorrhizal plant species) and Festuca rubra (a mycorrhizal plant species) were grown at defined atmospheric conditions differing in CO₂ concentration (350 and 700 ppm) for 3 years. During this period, C flow was assessed repeatedly (after 6 months, 1, 2, and 3 years) by ¹³C pulse‐chase experiments, and label was tracked through the rhizosphere bacterial, general fungal, and arbuscular mycorrhizal fungal (AMF) communities. Fatty acid biomarker analyses and RNA‐stable isotope probing (RNA‐SIP), in combination with real‐time PCR and PCR‐DGGE, were used to examine microbial community dynamics and abundance. Throughout the experiment the influence of elevated CO₂ was highly plant dependent, with the mycorrhizal plant exerting a greater influence on both bacterial and fungal communities. Biomarker data confirmed that rhizodeposited C was first processed by AMF and subsequently transferred to bacterial and fungal communities in the rhizosphere soil. Over the course of 3 years, elevated CO₂ caused a continuous increase in the ¹³C enrichment retained in AMF and an increasing delay in the transfer of C to the bacterial community. These results show that, not only do elevated atmospheric CO₂ conditions induce changes in rhizosphere C flow and dynamics but also continue to develop over multiple seasons, thereby affecting terrestrial ecosystems C utilization processes.     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Halving sunlight reveals no carbon limitation of aboveground biomass production in alpine grassland

In temperate alpine environments, the short growing season, low temperature and a slow nutrient cycle may restrict plant growth more than carbon (C) assimilation does. To test whether C is a limiting resource, we applied a shade gradient from ambient light to 44% (maximum shade) of incident photon flux density (PFD) in late successional, Carex curvula‐dominated alpine grassland at 2,580 m elevation in the Swiss central Alps for 3 years (2014–2016). Total aboveground biomass did not significantly decrease under reduced PFD, with a confidence interval ranging from +4% to ?15% biomass in maximum shade. Belowground biomass, of which more than 80% were fine roots, was significantly reduced by a mean of 17.9 ± 4.6% (±SE), corresponding to 228 g/m², in maximum shade in 2015 and 2016. This suggests reduced investments into water and nutrient acquisition according to the functional equilibrium concept. Specific leaf area (SLA) and maximum leaf length of the most abundant species increased with decreasing PFD. Foliar concentration of nonstructural carbohydrates (NSC) was reduced by 12.5 ± 4.3% under maximum shade (mean of eight tested species), while NSC concentration of belowground storage organs were unchanged in the four most abundant forbs. Furthermore, maximum shade lowered foliar δ¹³C by 1.56 ± 0.35‰ and increased foliar nitrogen concentrations per unit dry mass by 18.8 ± 4.1% across six species in 2015. However, based on unit leaf area, N concentrations were lower in shade (effect of higher SLA). Thus, while we found typical morphological and physiological plant responses to lower light, shading did not considerably affect seasonal aboveground biomass production of this alpine plant community within a broad range of PFD. This suggests that C is not a growth‐limiting resource, matching the unresponsiveness to in situ CO₂ enrichment previously reported for this type of grassland.     

Response to CO2 enrichment of understory vegetation in the shade of forests

Responses of forest ecosystems to increased atmospheric CO₂ concentration have been studied in few free‐air CO₂ enrichment (FACE) experiments during last two decades. Most studies focused principally on the overstory trees with little attention given to understory vegetation. Despite its small contribution to total productivity of an ecosystem, understory vegetation plays an important role in predicting successional dynamics and future plant community composition. Thus, the response of understory vegetation in Pinus taeda plantation at the Duke Forest FACE site after 15–17 years of exposure to elevated CO₂, 6–13 of which with nitrogen (N) amendment, was examined. Aboveground biomass and density of the understory decreased across all treatments with increasing overstory leaf area index (LAI). However, the CO₂ and N treatments had no effect on aboveground biomass, tree density, community composition, and the fraction of shade‐tolerant species. The increases of overstory LAI (~28%) under elevated CO₂ resulted in a reduction of light available to the understory (~18%) sufficient to nullify the expected growth‐enhancing effect of elevated CO₂ on understory vegetation.