Rapid root closure after fire limits fine root responses to elevated atmospheric CO2 in a scrub oak ecosystem in central Florida, USA

Elevated atmospheric carbon dioxide (CO₂) often stimulates the growth of fine roots, yet there are few reports of responses of intact root systems to long‐term CO₂ exposure. We investigated the effects of elevated CO₂ on fine root growth using open top chambers in a scrub oak ecosystem at Kennedy Space Center, Florida for more than 7 years. CO₂ enrichment began immediately after a controlled burn, which simulated the natural disturbance that occurs in this system every 10–15 years. We hypothesized that (1) root abundance would increase in both treatments as the system recovered from fire; (2) elevated CO₂ would stimulate root growth; and (3) elevated CO₂ would alter root distribution. Minirhizotron tubes were used to measure fine root length density (mm cm^?2) every three months. During the first 2 years after fire recovery, fine root abundance increased in all treatments and elevated CO₂ significantly enhanced root abundance, causing a maximum stimulation of 181% after 20 months. The CO₂ stimulation was initially more pronounced in the top 10 cm and 38–49 cm below the soil surface. However, these responses completely disappeared during the third year of experimental treatment: elevated CO₂ had no effect on root abundance or on the depth distribution of fine roots during years 3–7. The results suggest that, within a few years following fire, fine roots in this scrub oak ecosystem reach closure, defined here as a dynamic equilibrium between production and mortality. These results further suggest that elevated CO₂ hastens root closure but does not affect maximum root abundance. Limitation of fine root growth by belowground resources – particularly nutrients in this nutrient‐poor soil – may explain the transient response to elevated CO₂.     

Growth,loss, and vertical distribution of Pinus radiata fine roots growing at ambient and elevated CO2 concentration

Increased below-ground carbon allocation in forest ecosystems is a likely consequence of rising atmospheric CO₂ concentration. If this results in changes to fine root growth, turnover and distribution long-term soil carbon cycling and storage could be altered. Bi-weekly measurements were made to determine the dynamics and distribution of fine roots (< 1 mm diameter) for Pinus radiata trees growing at ambient (350 μmol mol^–1) and elevated (650 μmol mol^–1) CO₂ concentration in large open-top chambers. Measurements were made using minirhizotrons installed horizontally at depths of 0.1, 0.3, 0.5 and 0.9 m. During the first year, at a depth of 0.3 m, the increase in relative growth rate of roots occurred 6 weeks earlier in the elevated CO₂ treatment and the maximum rate was reached 10 weeks earlier than for trees in the ambient treatment. After 2 years, cumulative fine root growth (P_t) was 36% greater for trees growing at elevated CO₂ than at ambient CO₂ concentration, although this difference was not significant. A model of root growth driven by daily soil temperature accounted for between 43 and 99% of root growth variability. Total root loss (L_t) was 9% in the ambient and 14% in the elevated CO₂ treatment, although this difference was not significant. Root loss was greatest at 0.3 m. In the first year, 62% of fine roots grown between mid-summer and late-autumn disappeared within a year in the elevated CO₂ treatment, but only 18% in the ambient CO₂ treatment (P < 0.01). An exponential model relating L_t to time accounted for between 74 and 99% of the variability. Root cohort half-lives were 951 d for the ambient and 333 d for the elevated treatment. Root length density decreased exponentially with depth in both treatments, but relatively more fine roots grown in the elevated CO₂ treatment tended to occur deeper in the soil profile.     

Effects of elevated atmospheric CO2 on root decomposition in a scrub oak ecosystem

The effects of elevated atmospheric CO₂ on fine root decomposition over a 828‐day period were investigated using open top chambers with both ambient and elevated (700 ppm) CO₂ treatments in an oak–palmetto scrub ecosystem at Kennedy Space Center, Florida. Carbon dioxide enrichment of the chambers began 15 May 1996. The experiment included roots grown in ambient and elevated carbon dioxide. Vertical litterbags installed in September 1996 in each elevated and ambient chamber incubated from December 1996 to December 1998 showed no significant treatment effect on fine root or rhizome mass loss. Initial fine root percentage mass loss varied from 10.3% to 13.5% after three months; 55.5% to 38.3% of original mass had been lost after 828 days. A period of nitrogen immobilization occurred in both fine roots and rhizomes in the elevated CO₂ incubation, which is a potential mechanism for nitrogen conservation for this system in an elevated CO₂ world .     

Elevated atmospheric CO2 increases fine root production, respiration, rhizosphere respiration and soil CO2 efflux in Scots pine seedlings

In this study, we investigated the impact of elevated atmospheric CO₂ (ambient + 350 μmol mol^–1) on fine root production and respiration in Scots pine (Pinus sylvestris L.) seedlings. After six months exposure to elevated CO₂, root production measured by root in-growth bags, showed significant increases in mean total root length and biomass, which were more than 100% greater compared to the ambient treatment. This increased root length may have lead to a more intensive soil exploration. Chemical analysis of the roots showed that the roots in the elevated treatment accumulated more starch and had a lower C/N-ratio. Specific root respiration rates were significantly higher in the elevated treatment and this was probably attributed to increased nitrogen concentrations in the roots. Rhizospheric respiration and soil CO₂ efflux were also enhanced in the elevated treatment. These results clearly indicate that under elevated atmospheric CO₂ root production and development in Scots pine seedlings is altered and respiratory carbon losses through the root system are increased.     

Root Biomass of Individual Species, and Root Size Characteristics After Five Years of CO2 Enrichment on Native Shortgrass Steppe

Information from field studies investigating the responses of roots to increasing atmospheric CO₂ is limited and somewhat inconsistent, due partly to the difficulty in studying root systems in situ. In this report, we present standing root biomass of species and root length and diameter after five years of CO₂ enrichment (∽720 μmol mol⁻¹) in large (16 m² ground area) open-top chambers placed over a native shortgrass steppe in Colorado, USA. Total root biomass in 100 cm long×20 cm wide×75 cm depth soil monoliths and root biomass of the three dominant grass species of the site were not significantly affected by elevated CO₂. Root biomass of Stipa comata in the 0–20 cm soil depth was nearly 100% greater in elevated vs. ambient CO₂ chambers, but this was not statistically significant (P=0.14). However, there was a significant 37% increase in fine root length under elevated CO₂ in the 0–10 cm soil depth layer. Other reports from this study suggest that the increase in fine roots is primarily from improved seedling recruitment of S. comata under elevated CO₂. Few treatment differences in root length or diameter were detected in lower 10 cm depth increments, to 80 cm. These results reflect the root status integrated over two wet, two dry and one normal precipitation years and approximately one complete cycle of root turn-over on the shortgrass steppe. We conclude that increasing atmospheric CO₂ will have only small effects on standing root biomass and root length and diameter of most shortgrasss steppe species. However, the potential increased competitive ability of Stipa comata, a low forage quality species, could alter the ecosystem from the current dominant, high forage quality species, Bouteloua gracilis. B. gracilis is very well adapted to the frequent droughts of the shortgrass steppe. Increased competitive ability of less desirable plant species under increasing atmospheric CO₂ will have large implications for long-term sustainability of grassland ecosystems.     

The influence of elevated atmospheric CO2 on fine root dynamics in an intact temperate forest

Root dynamics are important for plant, ecosystem and global carbon cycling. Changes in root dynamics caused by rising atmospheric CO₂ not only have the potential to moderate further CO₂ increases, but will likely affect forest function. We used FACE (Free‐Air CO₂ Enrichment) to expose three 30‐m diameter plots in a 13‐year‐old loblolly pine (Pinus taeda) forest to elevated (ambient + 200 µL L^?1) atmospheric CO₂. Three identical fully instrumented plots were implemented as controls (ambient air only). We quantified root dynamics from October 1998 to October 1999 using minirhizotrons. In spite of 16% greater root lengths and 24% more roots per minirhizotron tube, the effects of elevated atmospheric CO₂ on root lengths and numbers were not statistically significant. Similarly, production and mortality were also unaffected by the CO₂ treatment, even though annual root production and mortality were 26% and 46% greater in elevated compared to ambient CO₂ plots. Average diameters of live roots present at the shallowest soil depth were, however, significantly enhanced in CO₂‐enriched plots. Mortality decreased with increasing soil depth and the slopes of linear regression lines (mortality vs. depth) differed between elevated and ambient CO₂ treatments, reflecting the significant CO₂ by depth interaction. Relative root turnover (root flux/live root pool) was unchanged by exposure to elevated atmospheric CO₂. Results from this study suggest modest, if any, increases in ecosystem‐level root productivity in CO₂‐enriched environments.     

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.     

Elevated CO2 and O3 effects on fine-root survivorship in ponderosa pine mesocosms

Atmospheric carbon dioxide (CO₂) and ozone (O₃) concentrations are rising, which may have opposing effects on tree C balance and allocation to fine roots. More information is needed on interactive CO₂ and O₃ effects on roots, particularly fine-root life span, a critical demographic parameter and determinant of soil C and N pools and cycling rates. We conducted a study in which ponderosa pine (Pinus ponderosa) seedlings were exposed to two levels of CO₂ and O₃ in sun-lit controlled-environment mesocosms for 3 years. Minirhizotrons were used to monitor individual fine roots in three soil horizons every 28 days. Proportional hazards regression was used to analyze effects of CO₂, O₃, diameter, depth, and season of root initiation on fine-root survivorship. More fine roots were produced in the elevated CO₂ treatment than in ambient CO₂. Elevated CO₂, increasing root diameter, and increasing root depth all significantly increased fine-root survivorship and median life span. Life span was slightly, but not significantly, lower in elevated O₃, and increased O₃ did not reduce the effect of elevated CO₂. Median life spans varied from 140 to 448 days depending on the season of root initiation. These results indicate the potential for elevated CO₂ to increase the number of fine roots and their residence time in the soil, which is also affected by root diameter, root depth, and phenology.     

Effects of elevated CO2 on fine root dynamics in a Mojave Desert community: a FACE study

Fine roots (≤1 mm diameter) are critical in plant water and nutrient absorption, and it is important to understand how rising atmospheric CO₂ will affect them as part of terrestrial ecosystem responses to global change. This study's objective was to determine the effects of elevated CO₂ on production, mortality, and standing crops of fine root length over 2 years in a free‐air CO₂ enrichment (FACE) facility in the Mojave Desert of southern Nevada, USA. Three replicate 25 m diameter FACE rings were maintained at ambient (～370 μmol mol^?1) and elevated CO₂ (～550 μmol mol^?1) atmospheric concentrations. Twenty‐eight minirhizotron tubes were placed in each ring to sample three microsite locations: evergreen Larrea shrubs, drought‐deciduous Ambrosia shrubs, and along systematic community transects (primarily in shrub interspaces which account for ～85% of the area). Seasonal dynamics were similar for ambient and elevated CO₂: fine root production peaked in April–June, with peak standing crop occurring about 1 month later, and peak mortality occurring during the hot summer months, with higher values for all three measures in a wet year compared with a dry year. Fine root standing crop, production, and mortality were not significantly different between treatments except standing crop along community transects, where fine root length was significantly lower in elevated CO₂. Fine root turnover (annual cumulative mortality/mean standing crop) ranged from 2.33 to 3.17 year^?1, and was not significantly different among CO₂ treatments, except for community transect tubes where it was significantly lower for elevated CO₂. There were no differences in fine root responses to CO₂ between evergreen (Larrea) and drought‐deciduous (Ambrosia) shrubs. Combined with observations of increased leaf‐level water‐use efficiency and lack of soil moisture differences, these results suggest that under elevated CO₂ conditions, reduced root systems (compared with ambient CO₂) appear sufficient to provide resources for modest aboveground production increases across the community, but in more fertile shrub microsites, fine root systems of comparable size with those in ambient CO₂ were required to support the greater aboveground production increases. For community transects, development of the difference in fine root standing crops occurred primarily through lower stimulation of fine root production in the elevated CO₂ treatment during periods of high water availability.     

Seasonal soil‐surface carbon fluxes from the root systems of young Pinus radiata trees growing at ambient and elevated CO2 concentration

Elevated atmospheric CO₂ concentration may result in increased below‐ground carbon allocation by trees, thereby altering soil carbon cycling. Seasonal estimates of soil surface carbon flux were made to determine whether carbon losses from Pinus radiata trees growing at elevated CO₂ concentration were higher than those at ambient CO₂ concentration, and whether this was related to increased fine root growth. Monthly soil surface carbon flux density (f) measurements were made on plots with trees growing at ambient (350) and elevated (650 μmol mol^?1) CO₂ concentration in large open‐top chambers. Prior to planting the soil carbon concentration (0.1%) and f (0.28 μmol m^?2 s^?1 at 15 °C) were low. A function describing the radial pattern of f with distance from tree stems was used to estimate the annual carbon flux from tree plots. Seasonal estimates of fine root production were made from minirhizotrons and the radial distribution of roots compared with radial measurements of f. A one‐dimensional gas diffusion model was used to estimate f from soil CO₂ concentrations at four depths. For the second year of growth, the annual carbon flux from the plots was 1671 g y^?1 and 1895 g y^?1 at ambient and elevated CO₂ concentrations, respectively, although this was not a significant difference. Higher f at elevated CO₂ concentration was largely explained by increased fine root biomass. Fine root biomass and stem production were both positively related to f. Both root length density and f declined exponentially with distance from the stem, and had similar length scales. Diurnal changes in f were largely explained by changes in soil temperature at a depth of 0.05 m. Ignoring the change of f with increasing distance from tree stems when scaling to a unit ground area basis from measurements with individual trees could result in under‐ or overestimates of soil‐surface carbon fluxes, especially in young stands when fine roots are unevenly distributed.     

Fine root biomass estimates from minirhizotron imagery in a shrub ecosystem exposed to elevated CO2

Fine root biomass and C content are critical components in ecosystem C models, but they cannot be directly determined by minirhizotron techniques, and indirect methods involve estimating 3-dimensional values (biomass/ soil volume) from 2-dimensional measurements. To estimate biomass from minirhizotron data, a conversion factor for length to biomass must be developed, and assumptions regarding depth of view must be made. In a scrub-oak ecosystem in central Florida, USA, root length density (RLD) was monitored for 10 years in a CO₂ manipulation experiment using minirhizotron tubes. In the seventh year of the study, soil cores were removed from both ambient and elevated CO₂ chambers. Roots from those cores were used to determine specific root length values (m/g) that were applied to the long-term RLD data for an estimation of root biomass over 10 years of CO₂ manipulation. Root length and biomass estimated from minirhizotron data were comparable to determinations from soil cores, suggesting that the minirhizotron biomass model is valid. Biomass estimates from minirhizotrons indicate the <0.25 mm diameter roots accounted for nearly 95% of the total root length in 2002. The long-term trends for this smallest size class (<0.25 mm diameter) mirrored the RLD trends closely, particularly in relation to suspected root closure in this system. Elevated CO₂ did not significantly affect specific root length as determined by the soil cores. A significant treatment effect indicated smallest diameter fine roots (<0.25 mm) were greater under elevated CO₂ during the early years of the study and the largest (2–10 mm) had greater biomass under elevated CO₂ during the later years of the study. Overall, this method permits long-term analysis of the effects of elevated CO₂ on fine root biomass accumulation and provides essential information for carbon models.     

CO2 and N-fertilization effects on fine-root length, production, and mortality: a 4-year ponderosa pine study

We conducted a 4-year study of juvenile Pinus ponderosa fine root (≤2 mm) responses to atmospheric CO₂ and N-fertilization. Seedlings were grown in open-top chambers at three CO₂ levels (ambient, ambient+175 μmol/mol, ambient+350 μmol/mol) and three N-fertilization levels (0, 10, 20 g m⁻² year⁻¹). Length and width of individual roots were measured from minirhizotron video images bimonthly over 4 years starting when the seedlings were 1.5 years old. Neither CO₂ nor N-fertilization treatments affected the seasonal patterns of root production or mortality. Yearly values of fine-root length standing crop (m m⁻²), production (m m⁻² year⁻¹), and mortality (m m⁻² year⁻¹) were consistently higher in elevated CO₂ treatments throughout the study, except for mortality in the first year; however, the only statistically significant CO₂ effects were in the fine-root length standing crop (m m⁻²) in the second and third years, and production and mortality (m m⁻² year⁻¹) in the third year. Higher mortality (m m⁻² year⁻¹) in elevated CO₂ was due to greater standing crop rather than shorter life span, as fine roots lived longer in elevated CO₂. No significant N effects were noted for annual cumulative production, cumulative mortality, or mean standing crop. N availability did not significantly affect responses of fine-root standing crop, production, or mortality to elevated CO₂. Multi-year studies at all life stages of trees are important to characterize belowground responses to factors such as atmospheric CO₂ and N-fertilization. This study showed the potential for juvenile ponderosa pine to increase fine-root C pools and C fluxes through root mortality in response to elevated CO₂.     

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.     

NMR imaging of root water distribution in intact Vicia faba L plants in elevated atmospheric CO2

The effect of elevated atmospheric CO₂ on water distribution in the intact roots of Vicia faba L. bean seedlings grown in natural soil was studied noninvasively with proton (¹H) nuclear magnetic resonance (NMR) imaging. Exposure of 24-d-old plants to atmospheric CO₂-enriched air at 650 cm³ m^?3 produced significant increases in water imaged in upper roots, hypogeal cotyledons and lower stems in response to a short-term drying-stress cycle. Above ground, drying produced negligible stem shrinkage and stomatal resistance was unchanged. In contrast, the same drying cycle caused significant depletion of water imaged in the same upper root structures in control plants subject to ambient CO₂ (350 m³ m^?3), and stem shrinkage and increased stomatal resistance. The results suggest that inhibition of transpiration caused by elevated CO₂ does not necessarily result in attenuation of water transport from lower root structures. Inhibition of water loss from upper roots and lower stem in elevated CO₂ environments may be a mitigating factor in assessing deleterious effects of greenhouse changes on crops during periods of dry climate.     

Effects of elevated atmospheric CO2 on fine root production and activity in an intact temperate forest ecosystem

We investigated the effects of elevated atmospheric CO₂ concentrations (ambient + 200 ppm) on fine root production and soil carbon dynamics in a loblolly pine (Pinus taeda) forest subject to free‐air CO₂ enrichment (FACE) near Durham, NC (USA). Live fine root mass (LFR) showed less seasonal variation than dead fine root mass (DFR), which was correlated with seasonal changes in soil moisture and soil temperature. LFR mass increased significantly (by 86%) in the elevated CO₂ treatment, with an increment of 37 g(dry weight) m^?2 above the control plots after two years of CO₂ fumigation. There was no long‐term increment in DFR associated with elevated CO₂, but significant seasonal accumulations of DFR mass occurred during the summer of the second year of fumigation. Overall, root net primary production (RNPP) was not significantly different, but annual carbon inputs were 21.7 gC m^?2 y^?1 (68%) higher in the elevated CO₂ treatment compared to controls. Specific root respiration was not altered by the CO₂ treatment during most of the year; however, it was significantly higher by 21% and 13% in September 1997 and May 1998, respectively, in elevated CO₂. We did not find statistically significant differences in the C/N ratio of the root tissue, root decomposition or phosphatase activity in soil and roots associated with the treatment. Our data show that the early response of a loblolly pine forest ecosystem subject to CO₂ enrichment is an increase in its fine root population and a trend towards higher total RNPP after two years of CO₂ fumigation.     

Root dynamics and demography in shortgrass steppe under elevated CO2, and comments on minirhizotron methodology

The dynamics and demography of roots were followed for 5 years that spanned wet and drought periods in native, semiarid shortgrass steppe grassland exposed to ambient and elevated atmospheric CO₂ treatments. Elevated compared with ambient CO₂ concentrations resulted in greater root‐length growth (+52%), root‐length losses (+37%), and total pool sizes (+41%). The greater standing pool of roots under elevated compared with ambient CO₂ was because of the greater number of roots (+35%), not because individuals were longer. Loss rates increased relatively less than growth rates because life spans were longer (+41%). The diameter of roots was larger under elevated compared with ambient CO₂ only in the upper soil profile. Elevated CO₂ affected root architecture through increased branching. Growth‐to‐loss ratio regressions to time of equilibrium indicate very long turnover times of 5.8, 7.0, and 5.3 years for control, ambient, and elevated CO₂, respectively. Production was greater under elevated compared with ambient CO₂ both below‐ and aboveground, and the above‐ to belowground ratios did not differ between treatments. However, estimates of belowground production differed among methods of calculation using minirhizotron data, as well as between minirhizotron and root‐ingrowth methods. Users of minirhizotrons may need to consider equilibration in terms of both new growth and disappearance, rather than just growth. Large temporal pulses of root initiation and termination rates of entire individuals were observed (analogous to birth–death rates), and precipitation explained more of the variance in root initiation than termination. There was a dampening of the pulsing in root initiation and termination under elevated CO₂ during both wet and dry periods, which may be because of conservation of soil water reducing the suddenness of wet pulses and duration and severity of dry pulses. However, a very low degree of synchrony was observed between growth and disappearance (production and decomposition).     

Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO2

To determine the long-term impact of elevated CO₂ on primary production of native tallgrass prairie, we compared the responses of tallgrass prairie at ambient and twice-ambient atmospheric CO₂ levels over an 8-year period. Plots in open-top chambers (4.5 m diameter) were exposed continuously (24 h) to ambient and elevated CO₂ from early April to late October each year. Unchambered plots were monitored also. Above-ground peak biomass was determined by clipping each year in early August, and root growth was estimated by harvesting roots from root ingrowth bags. Plant community composition was censused each year in early June. In the last 2 years of the study, subplots were clipped on 1 June or 1 July, and regrowth was harvested on 1 October. Volumetric soil water content of the 0–100 cm soil layer was determined using neutron scattering, and was generally higher in elevated CO₂ plots than ambient. Peak above-ground biomass was greater on elevated CO₂ plots than ambient CO₂ plots with or without chambers during years with significant plant water stress. Above-ground regrowth biomass was greater under elevated CO₂ than under ambient CO₂ in a year with late-season water stress, but did not differ in a wetter year. Root ingrowth biomass was also greater in elevated CO₂ plots than ambient CO₂ plots when water stress occurred during the growing season. The basal cover and relative amount of warm-season perennial grasses (C4) in the stand changed little during the 8-year period, but basal cover and relative amount of cool-season perennial grasses (C3) in the stand declined in the elevated CO₂ plots and in ambient CO₂ plots with chambers. Forbs (C3) and members of the Cyperaceae (C3) increased in basal cover and relative amount in the stand at elevated compared to ambient CO₂. Greater biomass production under elevated CO₂ in C4-dominated grasslands may lead to a greater carbon sequestration by those ecosystems and reduce peak atmospheric CO₂ concentrations in the future.     

Fine root turnover of Japanese white birch (<Emphasis Type="Italic">Betula platyphylla</Emphasis> var. <Emphasis Type="Italic">japonica</Emphasis>) grown under elevated CO<Subscript>2</Subscript> in northern Japan

Key message

Elevated CO ₂ reduced fine root dynamics (production and turnover) of white birch seedlings, especially grown in volcanic ash soil compared with brown forest soil.

Abstract

Increased atmospheric CO₂ usually enhances photosynthetic ability and growth of trees. To understand how increased CO₂ affects below-ground part of trees under varied soil condition, we investigated the responses of the fine root (diameter <2 mm) dynamics of Japanese white birch (Betula platyphylla var. japonica) which was planted in 2010. The three-year-old birch seedlings were grown in four experimental treatments comprising two levels of CO₂, i.e., ambient: 380–390 and elevated: 500 μmol mol^?1, in combination with two kinds of soil: brown forest (BF) soil and volcanic ash (VA) soil which has few nutrients. The growth and turnover of fine roots were measured for 3 years (2011–2013) using the Mini-rhizotron. In the first observation year, live fine root length (standing crop) in BF soil was not affected by CO₂ treatment, but it was reduced by the elevated CO₂ from the second observation year. In VA soil, live fine root length was reduced by elevated CO₂ for all 3 years. Fine root turnover tended to decrease under elevated CO₂ compared with ambient in both soil types during the first and second observation years. Turnover of fine root production and mortality was also affected by the two factors, elevated CO₂ and different soil types. Median longevity of fine root increased under elevated CO₂, especially in VA soil at the beginning, and a shorter fine root lifespan appeared after 2 years of observation (2011–2012). These results suggest that elevated CO₂ does not consistently stimulate fine root turnover, particularly during the plant seedlings stage, as it may depend on the costs and benefits of constructing and retaining roots. Therefore, despite the other uncontrollable environment factors, carbon sequestration to the root system may be varied by CO₂ treatment period, soil type and plant age.

     

Long-term effects of elevated atmospheric CO2 on below-ground biomass and transformations to soil organic matter in grassland

We determined the effects of elevated [CO₂] on the quantity and quality of below-ground biomass and several soil organic matter pools at the conclusion of an eight-year CO₂ enrichment experiment on native tallgrass prairie. Plots in open-top chambers were exposed continuously to ambient and twice-ambient [CO₂] from early April through late October of each year. Soil was sampled to a depth of 30 cm beneath and next to the crowns of C4 grasses in these plots and in unchambered plots. Elevated [CO₂] increased the standing crops of rhizomes (87%), coarse roots (46%), and fibrous roots (40%) but had no effect on root litter (mostly fine root fragments and sloughed cortex material >500 μm). Soil C and N stocks also increased under elevated [CO₂], with accumulations in the silt/clay fraction over twice that of particulate organic matter (POM; >53 μm). The mostly root-like, light POM (density ≤1.8 Mg m^-3) appeared to turn over more rapidly, while the more amorphous and rendered heavy POM (density >1.8 Mg m^-3) accumulated under elevated [CO₂]. Overall, rhizome and root C:N ratios were not greatly affected by CO₂ enrichment. However, elevated [CO₂] increased the C:N ratios of root litter and POM in the surface 5 cm and induced a small but significant increase in the C:N ratio of the silt/clay fraction to a depth of 15 cm. Our data suggest that 8 years of CO₂ enrichment may have affected elements of the N cycle (including mineralization, immobilization, and asymbiotic fixation) but that any changes in N dynamics were insufficient to prevent significant plant growth responses. This revised version was published online in June 2006 with corrections to the Cover Date.     

Leaf gas exchange and nitrogen dynamics of N2-fixing, field-grown Alnus glutinosa under elevated atmospheric CO2

Few studies have investigated the effects of elevated CO₂ on the physiology of symbiotic N₂-fixing trees. Tree species grown in low N soils at elevated CO₂ generally show a decline in photosynthetic capacity over time relative to ambient CO₂ controls. This negative adjustment may be due to a reallocation of leaf N away from the photosynthetic apparatus, allowing for more efficient use of limiting N. We investigated the effect of twice ambient CO₂ on net CO₂ assimilation (A), photosynthetic capacity, leaf dark respiration, and leaf N content of N2-fixing Alnus glutinosa (black alder) grown in field open top chambers in a low N soil for 160 d. At growth CO₂, A was always greater in elevated compared to ambient CO₂ plants. Late season A vs. internal leaf p(CO₂) response curves indicated no negative adjustment of photosynthesis in elevated CO₂ plants. Rather, elevated CO₂ plants had 16% greater maximum rate of CO₂ fixation by Rubisco. Leaf dark respiration was greater at elevated CO₂ on an area basis, but unaffected by CO₂ on a mass or N basis. In elevated CO₂ plants, leaf N content (μg N cm^?2) increased 50% between Julian Date 208 and 264. Leaf N content showed little seasonal change in ambient CO₂ plants. A single point acetylene reduction assay of detached, nodulated root segments indicated a 46% increase in specific nitrogenase activity in elevated compared to ambient CO₂ plants. Our results suggest that N₂-fixing trees will be able to maintain high A with minimal negative adjustment of photosynthetic capacity following prolonged exposure to elevated CO₂ on N-poor soils.