Independent, Interactive, and Species-Specific Responses of Leaf Litter Decomposition to Elevated CO2 and O3 in a Northern Hardwood Forest

The future capacity of forest ecosystems to sequester atmospheric carbon is likely to be influenced by CO₂-mediated shifts in nutrient cycling through changes in litter chemistry, and by interactions with pollutants like O₃. We evaluated the independent and interactive effects of elevated CO₂ (560 μl l⁻¹) and O₃ (55 nl l l⁻¹) on leaf litter decomposition in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) at the Aspen free air CO₂ enrichment (FACE) site (Wisconsin, USA). Fumigation treatments consisted of replicated ambient, +CO₂, +O₃, and +CO₂ + O₃ FACE rings. We followed mass loss and litter chemistry over 23 months, using reciprocally transplanted litterbags to separate substrate quality from environment effects. Aspen decayed more slowly than birch across all treatment conditions, and changes in decomposition dynamics of both species were driven by shifts in substrate quality rather than by fumigation environment. Aspen litter produced under elevated CO₂ decayed more slowly than litter produced under ambient CO₂, and this effect was exacerbated by elevated O₃. Similarly, birch litter produced under elevated CO₂ also decayed more slowly than litter produced under ambient CO₂. In contrast to results for aspen, however, elevated O₃ accelerated birch decay under ambient CO₂, but decelerated decay under enriched CO₂. Changes in decomposition rates (k-values) were due to CO₂- and O₃-mediated shifts in litter quality, particularly levels of carbohydrates, nitrogen, and tannins. These results suggest that in early-successional forests of the future, elevated concentrations of CO₂ will likely reduce leaf litter decomposition, although the magnitude of effect will vary among species and in response to interactions with tropospheric O₃.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Arbuscular mycorrhizae respond to plants exposed to elevated atmospheric CO2 as a function of soil depth

The importance of arbuscular mycorrhizae (AM) in plant and ecosystem responses to global changes, e.g. elevated atmospheric CO₂, is widely acknowledged. Frequently, increases in AM root colonization occur in response to increased CO₂, but also the lack of significant changes has been reported. The goal of this study was to test whether arbuscular mycorrhizae (root colonization and composition of root colonization) respond to plants grown in elevated CO₂ as a function of soil depth. We grew Bromus hordeaceus L. and Lotus wrangelianus Fischer & C. Meyer monocultures in large pots with a synthetic serpentine soil profile for 4 yr in an experiment, in which CO₂ concentration was crossed factorially with NPK fertilization. When analyzing root infection separately for topsoil (0–15 cm) and subsoil (15–45 cm), we found large (e.g., about 5-fold) increases of AM fungal root colonization in the subsoil in response to CO₂, but no significant changes in the corresponding topsoil of Bromus. Only the coarse endophyte AM fungi, not the fine endophyte AM fungi, were responsible for the observed increase in the bottom soil layer, indicating a depth-dependent shift in the AM community colonizing the roots, even at this coarse morphological level. Other response variables also had significant soil layer ^* CO₂ interaction terms. The subsoil response would have been hidden in an unstratified assessment of the total root system, since most of the root length was concentrated in the top soil layer. The increased presence of mycorrhizae in roots deeper in the soil should be considered in sampling protocols, as it may be indicative of changed patterns of nutrient acquisition and carbon sequestration.     

Moss Responses to Elevated CO2 and Variation in Hydrology in a Temperate Lowland Peatland

We studied the effects of elevated CO₂ (180–200 ppmv above ambient) on growth and chemistry of three moss species (Sphagnum palustre, S. recurvum and Polytrichum commune) in a lowland peatland in the Netherlands. Thereto, we conducted both a greenhouse experiment with both Sphagnum species and a field experiment with all three species using MiniFACE (Free Air CO₂ Enrichment) technology during 3 years. The greenhouse experiment showed that Sphagnum growth was stimulated by elevated CO₂ in the short term, but that in the longer term (≥1 year) growth was probably inhibited by low water tables and/or down-regulation of photosynthesis. In the field experiment, we did not find significant changes in moss abundance in response to elevated CO₂, although CO₂ enrichment appeared to reduce S. recurvum abundance. Both Sphagnum species showed stronger responses to spatial variation in hydrology than to increased atmospheric CO₂ concentrations. Polytrichum was insensitive to changes in hydrology. Apart from the confounding effects of hydrology, the relative lack of growth response of the moss species may also have been due to the relatively small increase in assimilated CO₂ as achieved by the experimentally added CO₂. We calculated that the added CO₂ contributed at most 32% to the carbon assimilation of the mosses, while our estimates based on stable C isotope data even suggest lower contributions for Sphagnum (24–27%). Chemical analyses of the mosses showed only small elevated CO₂ effects on living tissue N concentration and C/N ratio of the mosses, but the C/N ratio of Polytrichum was substantially lower than those of the Sphagnum species. Continuing expansion of Polytrichum at the expense of Sphagnum could reduce the C sink function of this lowland Sphagnum peatland, and similar ones elsewhere, as litter decomposition rates would probably be enhanced. Such a reduction in sink function would be driven mostly by increased atmospheric N deposition, water table regulation for agricultural purposes and land management to preserve the early successional stage (mowing, tree and shrub removal), since these anthropogenic factors will probably exert a greater control on competition between Polytrichum and Sphagnum than increased atmospheric CO₂ concentrations.     

Gas Exchange of Carrot Leaves in Response to Elevated CO2 Concentration

Short-term responses of four carrot (Daucus carota) cultivars: Cascade, Caro Choice (CC), Oranza, and Red Core Chantenay (RCC) to CO₂ concentrations (C _a) were studied in a controlled environment. Leaf net photosynthetic rate (P _N), intercellular CO₂ (C _i), stomatal conductance (g _s), and transpiration rate (E) were measured at C _a from 50 to 1 050 mol mol^–1. The cultivars responded similarly to C _a and did not differ in all the variables measured. The P _N increased with C _a until saturation at 650 mol mol^–1 (C _i= 350–400 mol mol^–1), thereafter P _N increased slightly. On average, increasing C _a from 350 to 650 and from 350 to 1 050 mol mol^–1 increased P _N by 43 and 52 %, respectively. The P _N vs. C _i curves were fitted to a non-rectangular hyperbola model. The cultivars did not differ in the parameters estimated from the model. Carboxylation efficiencies ranged from 68 to 91 mol m^–2 s^–1 and maximum P _N were 15.50, 13.52, 13.31, and 14.96 mol m^–2 s^–1 for Cascade, CC, Oranza, and RCC, respectively. Dark respiration rate varied from 2.80 mol m^–2 s^–1 for Oranza to 3.96 mol m^–2 s^–1 for Cascade and the CO₂ compensation concentration was between 42 and 46 mol mol^–1. The g _s and E increased to a peak at C _a= 350 mol mol^–1 and then decreased by 17 and 15 %, respectively when C _a was increased to 650 mol mol^–1. An increase from 350 to 1 050 mol mol^–1 reduced g _s and E by 53 and 47 %, respectively. Changes in g _s and P _N maintained the C _i:C _a ratio. The water use efficiency increased linearly with C _a due to increases in P _N in addition to the decline in E at high C _a. Hence CO₂ enrichment increases P _N and decreases g _s, and can improve carrot productivity and water conservation.     

The Photosynthetic Response to Elevated CO2 in High Altitude Potato Species (Solanum curtilobum)

Plants of Solanum curtilobum (from high altitude) and Solanum tuberosum (from low altitude) were grown in open-top chambers in a greenhouse at either ambient (AC, 360 µmol mol^–1) or ca. twice ambient (EC, 720 µmol mol^–1) CO₂ concentrations for 30 d. CO₂ treatments started at the reproductive stage of the plants. There were similar patterns in the physiological response to CO₂ enrichment in the two species. Stomatal conductance was reduced by 59 % in S. tuberosum and by 55 % in S. curtilobum, but such a reduction did not limit the net photosynthetic rate (P _N), which was increased by approximately 56 % in S. curtilobum and 53 % in S. tuberosum. The transpiration rate was reduced by 16 % in both potato species while instantaneous transpiration efficiency increased by 80 % in S. tuberosum and 90 % in S. curtilobum. Plants grown under EC showed 36 and 66 % increment in total dry biomass, whereas yields (dry mass of tubers) were increased by 40 and 85 % in S. tuberosum and S. curtilobum, respectively. EC promoted productivity by increasing P _N. Thus S. tuberosum, cultivated around the world at low altitudes, and S. curtilobum, endemic of the highland Andes, respond positively to EC during the tuberisation stage.     

Influence of Elevated CO2 on the Fungal Community in a Coastal Scrub Oak Forest Soil Investigated with Terminal-Restriction Fragment Length Polymorphism Analysis

Sixteen open-top chambers (diameter, 3.66 m) were established in a scrub oak habitat in central Florida where vegetation was removed in a planned burn prior to chamber installation. Eight control chambers have been continuously exposed to ambient air and eight have been continuously exposed to elevated CO₂ at twice-ambient concentration (~700 ppm) for 5 years. Soil cores were collected from each chamber to examine the influence of elevated atmospheric CO₂ on the fungal community in different soil fractions. Each soil sample was physically fractionated into bulk soil, rhizosphere soil, and roots for separate analyses. Changes in relative fungal biomass were estimated by the ergosterol technique. In the bulk soil and root fractions, a significantly increased level of ergosterol was detected in the elevated CO₂ treatments relative to ambient controls. Fungal community composition was determined by terminal-restriction fragment length polymorphism (T-RFLP) analysis of the internal transcribed spacer (ITS) region. The specificities of different ITS primer sets were evaluated against plant and fungal species isolated from the experimental site. Changes in community composition were assessed by principal component analyses of T-RFLP profiles resolved by capillary electrophoresis. Fungal species richness, defined by the total number of terminal restriction fragments, was not significantly affected by either CO₂ treatment or soil fraction.     

Controlled Environment Chambers for Investigating Tree Response to Elevated CO2 and Temperature Under Boreal Conditions

A closed CO₂ and temperature-controlled, long-term chamber system has been developed and set up in a typical boreal forest of Scots pine (Pinus sylvestris L.) near the Mekrijärvi Research Station (62°47N, 30°58E, 145 m above sea level) belonging to the University of Joensuu, Finland. The main objectives of the experiment were to provide a means of assessing the medium to long-term effects of elevated atmospheric CO₂ concentration (EC) and temperature (ET) on photosynthesis, respiration, growth, and biomass at the whole-tree level and to measure instantaneous whole-system CO₂ exchange. The system consists of 16 chambers with individual facilities for controlling CO₂ concentration, temperature, and the combination of the two. The chambers can provide a wide variety of climatic conditions that are similar to natural regimes. In this experiment the target CO₂ concentration in the EC chambers was set at a fixed constant of 700 µmol mol^–1 and the target air temperature in the ET chambers to track the ambient temperature but with a specified addition. Chamber performance was assessed on the base of recordings covering three consecutive years. The CO₂ and temperature control in these closed chambers was in general accurate and reliable. CO₂ concentration in the EC chambers was within 600–725 µmol mol^–1 for 90 % of the exposure time during the "growing-season" (15 April – 15 September) and 625–725 µmol mol^–1 for 88 % of the time in the "off-season" (16 September – 14 April), while temperatures in the chambers were within ±2.0 °C of the ambient or target temperature in the "growing season" and within ±3.0 °C in the "off season". There were still some significant chamber effects. Solar radiation in the chambers was reduced by 50–60 % for 82 % of the time in the "growing season" and 55–65 % for 78 % of the time in the "off season", and the relative humidity of the air was increased by 5–10 % for 72 % of the time in the "growing season" and 2–12 % for 91 % of the time in the "off season". The crown architecture and main phenophase of the trees were not modified significantly by enclosure in the chambers, but some physiological parameters changed significantly, e.g., the radiant energy-saturated photosynthesis rate, transpiration rate, maximum photochemical efficiency of photosystem 2, and chlorophyll content.     

Chlorophyll a Fluorescence Response of Norway Spruce Needles to the Long-Term Effect of Elevated CO2 in Relation to Their Position Within the Canopy

The long-term impact of elevated CO₂ concentration on photosynthetic activity of sun-exposed (E) versus shaded (S) foliage was investigated in a Picea abies stand (age 12 years) after three years of cultivation in adjustable-lamella-domes (ALD). One ALD is supplied with either ambient air [ca. 350 µmol(CO₂) mol^–1; AC-variant) and the second with elevated CO₂ concentration [ambient plus 350 µmol(CO₂) mol^–1; EC-variant). The pronounced vertical profile of the photosynthetically active radiation (PAR) led to the typical differentiation of the photosynthetic apparatus between the S- and E-needles in the AC-variant estimated from the irradiance-responses of various parameters of the room temperature chlorophyll (Chl) a fluorescence parameters. Namely, electron transport rate (ETR), photochemical efficiency of photosystem 2, PS2 (_PS2), irradiance-saturated values of non-photochemical quenching of minimum (SV₀) and maximum (NPQ) fluorescence levels, and photochemical fluorescence quenching (q_p) at higher irradiances were all significantly higher for E-needles as compared with the S-ones. The prolonged exposure to EC did not cause any stimulation of ETR for the E-needles but a strongly positive effect of EC on ETR was observed for the S-needles resulting in more than doubled ETR capacity in comparison with S-needles from the AC-variant. For the E-needles in EC-variant a slightly steeper reduction of the _PS2 and q_p occurred with the increasing irradiance as compared to the E-needles of AC-variant. On the contrary, the S-needles in EC variant revealed a significantly greater capacity to maintain a high _PS2 at irradiances lower than 200 µmol m^–2 s^–1 and to prevent the over-reduction of the PS2 reaction centres. Moreover, compared to the AC-variant the relation between SV₀ and NPQ exhibited a strong decrease (up to 72 %) of the SV₀-NPQ slope for the E-needles and an increase (up to 76 %) of this value for the S-needles. Hence the E- and S-foliage responded differently to the long-term impact of EC. Moreover, this exposure was responsible for the smoothing of the PAR utilisation vertical gradient in PS2 photochemical and non-photochemical reactions within the canopy.     

Response of Forest Trees to Increased Atmospheric CO 2

The CO ₂ fertilization hypothesis stipulates that rising atmospheric CO ₂ has a positive effect on tree growth due to increasing availability of carbon. The objective of this paper is to compare the recent literature related to both field CO ₂ -enriched experiments with trees and empirical dendrochronological studies detecting CO ₂ fertilization effects in tree-rings. This will allow evaluation of tree growth responses to atmospheric CO ₂ enrichment by combining evidence from both ecophysiology and tree-ring research. Based on considerable experimental evidence of direct CO ₂ fertilization effect (increased photosynthesis, water use efficiency, and above- and belowground biomass), and predications from the interactions of enriched CO ₂ with temperature, nitrogen and drought, we propose that warm, moderately drought-stressed ecosystems with an ample nitrogen supply might be the most CO ₂ responsive ecosystems. Empirical tree-ring studies took the following three viewpoints on detecting CO ₂ fertilization effect in tree-rings: 1) finding evidence of CO ₂ fertilization effect in tree-rings, 2) attributing growth enhancement to favorable climate rather than atmospheric CO ₂ enrichment, and 3) considering that tree growth enhancement might be caused by synergistic effects of several factors such as favorable climate change, CO ₂ fertilization, and anthropogenic atmospheric deposition (e.g., nitrogen). At temperature-limiting sites such as high elevations, nonfindings of CO ₂ fertilization evidence could be ascribed to the following possibilities: 1) cold temperatures, a short season of cambial division, and nitrogen deficiency that preclude a direct CO ₂ response, 2) old trees past half of their maximum life expectancy and consequently only a small increase in biomass increment due to CO ₂ fertilization effect might be diminished, 3) the elimination of age/size-related trends by statistical detrending of tree-ring series that might remove some long-term CO ₂ -related trends in tree-rings, and 4) carbon partitioning and growth within a plant that is species-specific. Our review supports the atmospheric CO ₂ fertilization effect hypothesis, at least in trees growing in semi-arid or arid conditions because the drought-stressed trees could benefit from increased water use efficiency to enhance growth.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.     

Vascular Plant Responses to Elevated CO2 in a Temperate Lowland Sphagnum Peatland

Vascular plant responses to experimental enrichment with atmospheric carbon dioxide (CO₂), using MINIFACE technology, were studied in a Dutch lowland peatland dominated by Sphagnum and Phragmites for 3 years. We hypothesized that vascular plant carbon would accumulate in this peatland in response to CO₂ enrichment owing to increased productivity of the predominant species and poorer quality (higher C/N ratios) and consequently lower decomposability of the leaf litter of these species. Carbon isotope signatures demonstrated that the extra 180 ppmv CO₂ in enriched plots had been incorporated into vegetation biomass accordingly. However, on the CO₂ sequestration side of the ecosystem carbon budget, there were neither any significant responses of total aboveground abundance of vascular plants, nor of any of the individual species. On the CO₂ release side of the carbon budget (decomposition pathway), litter quantity did not differ between ambient and CO₂ treatments, while the changes in litter quality (N and P concentration, C/N and C/P ratio) were marginal and inconsistent. It appeared therefore that the afterlife effects of significant CO₂-induced changes in green-leaf chemistry (lower N and P concentrations, higher C/N and C/P) were partly offset by greater resorption of mobile carbohydrates from green leaves during senescence in CO₂-enriched plants. The decomposability of leaf litters of three predominant species from ambient and CO₂-enriched plots, as measured in a laboratory litter respiration assay, showed no differences. The relatively short time period, environmental spatial heterogeneity and small plot sizes might explain part of the lack of CO₂ response. When our results are combined with those from other Sphagnum peatland studies, the common pattern emerges that the vascular vegetation in these ecosystems is genuinely resistant to CO₂-induced change. On decadal time-scales, water management and its effects on peatland hydrology, N deposition from anthropogenic sources and land management regimes that arrest the early successional phase (mowing, tree and shrub removal), may have a greater impact on the vascular plant species composition, carbon balance and functioning of lowland Sphagnum–Phragmites reedlands than increasing CO₂ concentrations in the atmosphere.     

Vertical distribution and seasonal changes of fine and coarse root mass in Pinus kesiya Royle Ex.Gordon forest of three different ages

Seasonal changes and vertical distribution of fine (< 2 mm diameter) and coarse (2-10 mm diameter) root mass of Pinus kesiya and fine root and rhizome mass of herbaceous species, and root production were studied in the 6-, 15- and 23-year old Pinus kesiya forest stands at Shillong, in the Meghalaya state of north-east India. Maximum fine and coarse root mass of P. kesiya, and fine root and rhizome mass of the ground vegetation were recorded during the rainy season. The contribution of the tree fine roots in 0-10 cm soil layer declined from 51% in the 6-year old stand to about 33% in the older stands. The major proportion (63-88%) of herbaceous fine root and rhizome mass was concentrated in this soil layer in all the three stands. The majority (36-57%) of tree coarse roots were present in the 10-20 cm layer in all the stands. The biomass and necromass values in the case of fine roots were more or less equal in a given stand, but the coarse roots had 5 to 9 times more live than the dead mass. The proportion of herbaceous fine root mass to the total fine root mass declined from 54% in the 6-year old stand to 30-32% in the 15- and 23-year old stands. The mean total fine root mass (pine + herbaceous species) decreased from 417 g m^–2 in the 6-year old stand to 302 in 15-year and 322 g m^–2 in the 23-year old stand. Annual fine root production showed a marked decrease from 1055 g m^–2 in the 6-year old stand to 743 g m^–2 in the 23-year old stand, but coarse root production increased from 169 g m^–2 in the 6-year to 466 g m^–2  in the 23-year old stand; the total root production thus remained approximately constant.     

Fine root growth phenology,production, and turnover in a northern hardwood forest ecosystem

A large part of the nutrient flux in deciduous forests is through fine root turnover, yet this process is seldom measured. As part of a nutrient cycling study, fine root dynamics were studied for two years at Huntington Forest in the Adirondack Mountain region of New York, USA. Root growth phenology was characterized using field rhizotrons, three methods were used to estimate fine root production, two methods were used to estimate fine root mortality, and decomposition was estimated using the buried bag technique. During both 1986 and 1987, fine root elongation began in early April, peaked during July and August, and nearly ceased by mid-October. Mean fine root ( 3 mm diameter) biomass in the surface 28-cm was 2.5 t ha^–1 and necromass was 2.9 t ha^–1. Annual decomposition rates ranged from 17 to 30% beneath the litter and 27 to 52% at a depth of 10 cm. Depending on the method used for estimation, fine root production ranged from 2.0 to 2.9 t ha^–1, mortality ranged from 1.8 to 3.7 t ha^–1 yr^–1, and decomposition was 0.9 t ha^–1 yr^–1. Thus, turnover ranged from 0.8 to 1.2 yr^–1. The nutrients that cycled through fine roots annually were 4.5–6.1 kg Ca, 1.1–1.4 kg Mg, 0.3–0.4 kg K, 1.2–1.7 kg P, 20.3–27.3 kg N, and 1.8–2.4 kg S ha^–1. Fine root turnover was less important than leaf litterfall in the cycling of Ca and Mg and was similar to leaf litterfall in the amount of N, P, K and S cycled.     

Response of ectomycorrhizalPinus banksiana andPicea glauca to heavy metals in soil

Pinus banksiana andPicea glauca inoculated or not with the ectomycorrhizal fungusSuillus luteus were grown in a sandy loam soil containing a range of Cd, Cu, Ni, Pb and Zn concentrations. Ectomycorrhizal colonization rates were significantly reduced on Pinus and Picea seedlings by the heavy metals, particularly Cd and Ni. Needle tissue metal concentrations were lower in ectomycorrhizal seedlings at low soil metal concentrations. However, at higher soil concentrations, heavy metal concentrations of needle tissue were similar in ectomycorrhizal and nonmycorrhizal plants. The growth of nonmycorrhizal seedlings exposed to heavy metals was reduced compared to those inoculated withSuillus luteus. Apparently ectomycorrhizal colonization can protect Pinus and Picea seedlings from heavy metal toxicity at low or intermediate soil concentrations of Cd, Cu, Ni, Pb and Zn.     

Estimation of fine root production, mortality and turnover with Minirhizotron in Larix gmelinii and Fraxinus mandshurica plantations

Fine root turnover is a major pathway for carbon and nutrient cycling in forest ecosystems. However, to estimate fine root turnover, it is important to first understand the fine root dynamic processes associated with soil resource availability and climate factors. The objectives of this study were: (1) to examine patterns of fine root production and mortality in different seasons and soil depths in the Larix gmelinii and Fraxinus mandshurica plantations, (2) to analyze the correlation of fine root production and mortality with environmental factors such as air temperature, precipitation, soil temperature and available nitrogen, and (3) to estimate fine root turnover. We installed 36 Minirhizotron tubes in six mono-specific plots of each species in September 2003 in the Mao’ershan Experimental Forest Station. Minirhizotron sampling was conducted every two weeks from April 2004 to April 2005. We calculated the average fine root length, annual fine root length production and mortality using image data of Minirhizotrons, and estimated fine root turnover using three approaches. Results show that the average growth rate and mortality rate in L. melinii were markedly smaller than in F. mandshurica, and were highest in the surface soil and lowest at the bottom among all the four soil layers. The annual fine root production and mortality in F. mandshurica were significantly higher than in L. gmelinii. The fine root production in spring and summer accounted for 41.7% and 39.7% of the total annual production in F. mandshurica and 24.0% and 51.2% in L. gmelinii. The majority of fine root mortality occurred in spring and summer for F. mandshurica and in summer and autumn for L. gmelinii. The turnover rate was 3.1 a⁻¹ for L. gmelinii and 2.7 a⁻¹ for F. mandshurica. Multiple regression analysis indicates that climate and soil resource factors together could explain 80% of the variations of the fine root seasonal growth and 95% of the seasonal mortality. In conclusion, fine root production and mortality in L. gmelinii and F. mandshurica have different patterns in different seasons and at different soil depths. Air temperature, precipitation, soil temperature and soil available nitrogen integratively control the dynamics of fine root production, mortality and turnover in both species. Transtlated from Journal of Plant Ecology, 2007, 31(2): 333–342 [译自: 植物生态学报]     

Glass Domes with Adjustable Windows: A Novel Technique for Exposing Juvenile Forest Stands to Elevated CO2 Concentration

We present a new technological approach for in situ investigation of long-term impacts of elevated CO₂ concentration (EC) on juvenile forests characterised by an intensive community level and canopy closure phase. Construction of the glass domes is based on the properties of earlier tested open-top chambers (OTCs). An air climatisation device together with an adjustable window system, that forms the shell cover of the domes, is able to keep the required [CO₂] in both time and spatial scales with the relatively small consumption of supplied CO₂. This is achieved by half-closing the windows on the windward side. We evidenced good coupling of treated trees to the atmosphere, including mutual interactions among trees. The semi-open design of the domes moderates the problems of strong wind, humidity, and temperature gradients associated with OTCs. The frequency distributions of the environmental variations within the domes indicate that: air temperature is maintained within the ambient range ±1.0 °C for ca. 80 % of the time, and changes in the relative air humidity vary from –15 to 0 % for ca. 82 % of the time. The most important chamber effect is associated with the penetration of solar irradiance, which is reduced by 26 % compared to the open condition outside the domes. The dimensions of the domes are 10×10 m in length and 7 m high in the central part. The experiment was done in three identical stands of twelve-year-old Norway spruce trees. The 56 trees are planted at two different spacings to estimate the impacts of stand spatial structure in relation to EC.     

Increased Quantity and Quality of Coarse Soil Organic Matter Fraction at Elevated CO2 in a Grazed Grassland are a Consequence of Enhanced Root Growth Rate and Turnover

The aims of this study were to determine whether elevated atmospheric CO₂ concentration modifies plant organic matter (OM) fluxes to the soil and whether any change in the fluxes can modify soil OM accumulation. Measurements were made in a grazed temperate grassland after almost 4 years exposure to elevated atmospheric CO₂ (475 μl l^-1) using a Free Air CO₂ Enrichment (FACE) facility located in the North Island of New Zealand. Aboveground herbage biomass and leaf litter production were not altered by elevated CO₂ but root growth rate, as measured with the ingrowth core method, and root turnover were strongly stimulated by elevated CO₂ particularly at low soil moisture contents during summer. Consequently, significantly more plant material was returned to the soil under elevated CO₂ leading to an accumulation of coarse (> 1 mm) particulate organic matter (POM) but not of finer POM fractions. The accumulating POM exhibited a lower C/N ratio, which was attributed to the higher proportion of legumes in the pasture under elevated CO₂. Only small changes were detected in the size and activity of the soil microbial biomass in response to the POM accumulation, suggesting that higher organic substrate availability did not stimulate microbial growth and activity despite the apparent lower C/N ratio of accumulating POM. As a result, elevated CO₂ may well lead to an accumulation of OM in grazed grassland soil in the long term.     

Above- and Belowground Net Primary Production in a Temperate Mixed Deciduous Forest

Our current ability to detect and predict changes in forest ecosystem productivity is constrained by several limitations. These include a poor understanding of belowground productivity, the short duration of most analyses, and a need for greater examination of species- or community-specific variability in productivity studies. We quantified aboveground net primary productivity (ANPP) over 3 years (1999–2001), and both belowground NPP (BNPP) and total NPP over 2 years (2000–2001) in both mesic and xeric site community types of the mixed mesophytic forest of southeastern Kentucky to examine landscape variability in productivity and its relation with soil resource [water and nitrogen (N)] availability. Across sites, ANPP was significantly correlated with N availability (R² = 0.58, P = 0.028) while BNPP was best predicted by soil moisture content (R² = 0.72, P = 0.008). Because of these offsetting patterns, total NPP was unrelated to either soil resource. Interannual variability in growing season precipitation during the study resulted in a 50% decline in mesic site litter production, possibly due to a lag effect following a moderate drought year in 1999. As a result, ANPP in mesic sites declined 27% in 2000 compared to 1999, while xeric sites had no aboveground production differences related to precipitation variability. If global climate change produces more frequent occurrences of drought, then the response of mesic sites to prolonged moisture deficiency and the consequences of shifting carbon (C) allocation on C storage will become important questions.     

Effects of elevated soil solution Al concentrations on fine roots in a middle-aged Norway spruce (Picea abies (L.) Karst.) stand

Aluminium (Al), mobilized by acidic deposition, has been claimed to be a major threat to forest vitality. Fine root mortality, decreased root growth and reduced nutrient uptake have been observed in controlled laboratory experiments where roots of tree seedlings were exposed to elevated concentrations of Al. Yet, evidence for Al-induced root damage from forest stands is scarcely reported. Nevertheless, Al dissolved in soil water has received a key role in the critical load concept for forests. Here, we present effects of artificially elevated concentrations of Al in the soil solution on fine roots in a middle-aged stand of Norway spruce (Picea abies (L.) Karst.). Although the inorganic Al concentrations about 200 µM and Ca:Al ratio about 0.7 that were established in the soil solution within this experiment have been associated with reduction of root growth and root mortality for spruce seedlings in hydroponic studies, no acute damage on fine roots was observed. Three years of treatment did not cause visual root damage, nor were effects on fine root necromass observed. Fine root necromass made up about 10% of fine root biomass for all treatments. However, significantly lower molar Ca:Al and Mg:Al ratios in living and dead fine roots were found in the plots where Al concentrations were highest and ratios of Ca to Al in the soil solution were lowest. The lack of response on fine root biomass suggests that forest stands tolerate higher Al levels than results from laboratory experiments indicate. We conclude that effect studies in the laboratory have limited value for field conditions. The key role of Al toxicity, expressed as the Ca/Al ratio, in critical load calculations for forests may have to be reconsidered.