Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands

Fine root systems may respond to soil chemical conditions, but contrasting results have been obtained from field studies in non-manipulated forests with distinct soil chemical properties. We investigated biomass, necromass, live/dead ratios, morphology and nutrient concentrations of fine roots (<2 mm) in four mature Norway spruce (Picea abies [L.] Karst.) stands of south-east Germany, encompassing variations in soil chemical properties and climate. All stands were established on acidic soils (pH (CaCl₂) range 2.8–3.8 in the humus layer), two of the four stands had molar ratios in soil solution below 1 and one of the four stands had received a liming treatment 22 years before the study. Soil cores down to 40 cm mineral soil depth were taken in autumn and separated into four fractions: humus layer, 0–10 cm, 10–20 cm and 20–40 cm. We found no indications of negative effects of N availability on fine root properties despite large variations in inorganic N seepage fluxes (4–34 kg N ha⁻¹ yr⁻¹), suggesting that the variation in N deposition between 17 and 26 kg N ha⁻¹ yr⁻¹ does not affect the fine root system of Norway spruce. Fine root biomass was largest in the humus layer and increased with the amount of organic matter stored in the humus layer, indicating that the vertical pattern of fine roots is largely affected by the thickness of this horizon. Only two stands showed significant differences in fine root biomass of the mineral soil which can be explained by differences in soil chemical conditions. The stand with the lowest total biomass had the lowest Ca/Al ratio of 0.1 in seepage, however, Al, Ca, Mg and K concentrations of fine roots were not different among the stands. The Ca/Al ratio in seepage might be a less reliable stress parameter because another stand also had Ca/Al ratios in seepage far below the critical value of 1.0 without any signs of fine root damages. Large differences in the live/dead ratio were positively correlated with the Mn concentration of live fine roots from the mineral soil. This relationship was attributed to faster decay of dead fine roots because Mn is known as an essential element of lignin degrading enzymes. It is questionable if the live/dead ratio can be used as a vitality parameter of fine roots since both longevity of fine roots and decay of root litter may affect this parameter. Morphological properties were different in the humus layer of one stand that was limed in 1983, indicating that a single lime dose of 3–4 Mg ha⁻¹ has a long-lasting effect on fine root architecture of Norway spruce. Almost no differences were found in morphological properties in the mineral soil among the stands, but vertical patterns were apparently different. Two stands with high base saturation in the subsoil showed a vertical decrease in specific root length and specific root tip density whereas the other two stands showed an opposite pattern or no effect. Our results suggest that proliferation of fine roots increased with decreasing base saturation in the subsoil of Norway spruce stands.     

Ammonium,nitrate and dissolved organic nitrogen in seepage water as affected by compost amendment to European beech,Norway spruce,and Scots pine forests

Fertilization of nutrient-depleted and degraded forest soils may be required to sustain utilization of forests. In some European countries, the application of composts may now be an alternative to the application of inorganic fertilizers because commercial compost production has increased and compost quality has been improved. There is, however, concern that compost amendments may cause increased leaching of nitrogen, trace metals and toxic organic compounds to groundwater. The objective of this study was to assess the risk of ammonium (NH₄ ⁺), nitrate (NO₃ ^–) and dissolved organic nitrogen (DON) leaching following a single compost application to silty and sandy soils in mature beech (Fagus sylvatica L.), pine (Pinus silvestris L.) and spruce (Picea abies Karst.) forests at Solling and Unterlüß in Lower Saxony, Germany. Mature compost from separately collected organic household waste was applied to the soil surface at a rate of 6.3 kg m^–2 in the summer of 1997 and changes in NH₄ ⁺, NO₃ ^– and DON concentrations in throughfall and soil water at 10 and 100 cm soil depths were determined for 32 months. The spruce forests had the highest N inputs by throughfall water and the highest N outputs in both the control and compost plots compared with the pine and beech forests. Overall, the differences in total N outputs at 100 cm soil depth between the control and compost plots ranged between 0.3 and 11.2 g N m^–2 for the entire 32-month period. The major leaching of these amounts occurred during the first 17 months after compost amendments, but there was no significant difference in total N outputs (–0.2 to 1.8 g N m^–2) between the control and compost plots during the remaining 15 months. Most of the mineral soils acted as a significant sink for NO₃ ^– and DON as shown by a reduction of their outputs from 10 to 100 cm depth. Based on these results, we conclude that application of mature compost with high inorganic N contents could diminish the groundwater quality in the first months after the amendments. A partial, moderate application of mature compost with low inorganic N content to nutrient depleted forest soils can minimize the risk of NO₃ ^– leaching.     

Influence of Norway spruce seedlings on the nutrient availability in mineral soil and forest floor material

A greenhouse experiment was performed to evaluate the effect of Norway spruce (Picea abies (L.) Karst.) seedlings on net nutrient availability in five different growing media containing F- or H-layer and mineral soil originating from a haplic podzol in northern Sweden. The initial total amounts of eight nutrient elements (N, K, P, Ca, Mg, Mn, Fe, Zn) and exchangeable amounts of the same elements were analyzed in pots with or without spruce seedlings. In the planted pots seedling nutrient uptake was also estimated. After 26 weeks, higher net nutrient availability with seedlings was found in 25 out of the 40 (62%) growing media and nutrient element combinations. A positive seedling effect on net nutrient availability might be explained by rhizodeposition stimulating the soil microorganism activity and accelerating the weathering of minerals or by seedling roots promoting the nutrient providing processes through changes in soil chemical and physical properties. Nitrogen availability was primarily affected by what part of the forest floor the growing medium contained although the positive response to seedling presence was apparent. The positive net availability response of P, Ca, Mg, Mn, Fe and Zn to seedling presence was on the other hand relatively strong. In the case of P, K, and Zn the growing medium composition (if the F- and H-layer was pure or mixed with mineral soil) was also an important factor for the estimated net availability. Pure F-and H-layer provided greater P- and K-availability while the availability of Zn increased when mineral soil was added. The influence of growing plants ought to be considered when soil samples are used for assessing the nutrient availability.     

Effects of growth medium, nutrients, water, and aeration on mycorrhization and biomass allocation of greenhouse-grown interior Douglas-fir seedlings

Commercial nursery practices usually fail to promote mycorrhization of interior Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco] seedlings in British Columbia, which may account for their poor performance following planting in the field. We tested the effects of four nursery cultivation factors (nitrogen fertilization, phosphorus fertilization, watering, and soil aeration) and field soil addition on mycorrhization, survival, growth, and biomass allocation of interior Douglas-fir seedlings in a series of greenhouse experiments. Where field soil was added to the growing medium, mycorrhization and root/shoot ratios were maximized at lower levels of mineral nutrient application and aeration. Where field soil was not added, mycorrhization was negligible across all fertilization and aeration treatments, but root/shoot ratio was maximized at lower levels of mineral nutrients and the highest level of aeration. Regardless of whether field soil was added, intermediate levels of soil water resulted in the best mycorrhizal colonization and root/shoot ratios. However, field soil addition reduced seedling mortality at the two lowest water levels. A cluster analysis placed ectomycorrhizal morphotypes into three groups (Mycelium radicis-atrovirens Melin, Wilcoxina, and mixed) based on their treatment response, with all but two morphotypes in the mixed group whose abundance was maximized under conditions common to advanced seedling establishment. For maximal mycorrhization and root development of interior Douglas-fir seedlings, nurseries should minimize addition of nitrogen and phosphorus nutrients, maximize aeration, provide water at moderate rates, and, where possible, add small amounts of field soil to the growing medium.     

Monitoring uptake and contents of Mg,Ca and K in Norway spruce as influenced by pH and Al,using microprobe analysis and stable isotope labelling

In a model system using intact spruce trees (Picea abies [L.] Karst.) we followed the path of magnesium, calcium and potassium during uptake into the root and during long-range transport into the shoot, by multiple stable isotope labelling. The roots of two- and three-year-old spruce trees originating from soil culture were removed from the soil and, in part or in toto, exposed to labelling solutions containing the stable isotopes ²⁵Mg or ²⁶Mg, ⁴¹K and ⁴²Ca or ⁴⁴Ca. Optical-emission-spectroscopy (ICP-OES) of plant fractions and labelling solutions was combined with the quantitative analysis of stable isotope ratios in sections of shock frozen, cryosubstituted material using the laser-microprobe-mass-analyser (LAMMA). This combination allowed us to distinguish, both in bulk samples and on the cellular level between (i) the fraction of elements originally present in the plant before the start of the labelling, (ii) the material taken up from the labelling solution into the plant and (iii) any material released by the plant into the labelling solution.In single-root labelling experiments, roots of three-year-old spruce trees, grown in nursery soil, were exposed to various pH conditions. The exchange of Mg and Ca with the labelling solution was nearly 100% in the cell walls of the mycorrhized finest roots. This exchange was only slightly affected by a step down to pH 3.5. The absolute Mg and Ca content in the cell walls was moderately reduced by incubation at pH 3.5 and strongly reduced in the presence of Al at this pH. After a pH 3.5 and 2 mM Al treatment we found Al in the xylem cell walls and the cortex cell lumina at elevated concentrations. To analyse the combined effect of high Al and high proton concentrations on the long-range transport, we used a split-root system. The root mass of an intact two-year-old spruce tree, grown in mineral soil, was divided into even parts and both halves incubated in solutions with two sets of different stable isotopes of Mg and Ca (side A: no Al, ²⁵Mg and ⁴²Ca; side B: +Al, ²⁶Mg and ⁴⁴Ca) and ⁴¹K on both sides. We observed a large uptake of Mg, Ca and K into the plant and a pronounced release. The net uptake of all three elements was lower from the Al-doted solution. In cross-sections of the apical shoot we found after seven-day labelling period about 60–70% of the Mg and Ca and 30% of the K content in the xylem cell walls originating from both labelling solutions. The clear majority of the Mg and Ca label originated from the Al-doted side.     

Effect of exposure to fluoride,nitrogen compounds and SO2 on the numbers of spruce shoot aphids on Norway spruce seedlings

Summary Development of spruce shoot aphid (Cinara pilicornis Hartig) populations was monitored in natural and artificial infestations of Norway spruce (Picea abies Karst.) seedlings, exposed to air pollutants in an experimental field. The pollutants, applied both singly and in mixtures, were gaseous sulphur dioxide, NaF (30 mg l^-1 F) and Ca(NO₃)₂ or (NH₄)₂SO₄ in aqueous solutions (200 mg l^-1 N). Aphid numbers on 10 seedlings in each treatment and two control plots were counted at 2-week intervals. At the beginning of the experiment aphid numbers did not differ between treatments. Aphid populations peaked in late June and early July. All the pollutants and their combinations significantly increased the numbers of aphids per seedling. Four apterous females were transferred to spruce seedlings which were growing in containers in the same plots. After 4–5 weeks aphid numbers were significantly higher in the fluoride treatment and in the combined treatment of fluoride, nitrogen and SO₂. The pollution treatments did not have a significant effect on shoot growth. Concentrations of F and S in needles were higher in treatments involving these pollutants. There were no significant differences in the concentrations of free amino acids in shoot stems between control and fluoride treatment. However, the relatively low concentration of arginine in the F treatment at the end of the growing season might indicate disturbances in the nitrogen metabolism of spruce seedlings.     

Effect of initial fertilisation on biomass and nutrient contentof Norway spruce and Douglas-fir plantations at the same site

Norway spruce (NS) and Douglas-fir (DF) are among the main species used for production forestry in France. In low-elevation mountains and under-acidic conditions, they often occupy the same ecological situations. It is therefore of paramount interest to have a good understanding of how the two species behave under similar conditions and how they react to site improvement by fertilisation. The study stands are part of an experimental stand located in the estate forest of Breuil-Chenue in the Morvan (east central part of France). Its aim is to compare the impact of change in species on ecosystem functions. Destructive sampling of 10 trees per stand, distributed over the whole spectrum of inventoried classes of circumference at breast height (c _1.30), was carried out within four stands, e.g., fertilised and control (non-fertilised) NS; fertilised and control (non-fertilised) DF. Allometric relationships between c _1.30 and biomass or nutrient content per tree compartment were calculated. These equations were applied to the stand inventory for quantifying stand biomass and nutrient content on a hectare basis. The standard deviations of results were estimated using Monte-Carlo simulations. Specific emphasis was given to explain the origin of differences observed between species and treatments, i.e., changes in carbon allocation leading to specific allometric relationships, changes in stand structure (tree size distributions) and changes in stand density due to mortality.DF was more productive than NS (+28% for total tree biomass, +50% for ligneous biomass and +53% for stem wood). Both NS and DF were affected by fertilisation but in the case of NS, effects on the crown_c _1.30 relationship and on average tree growth were predominant while in the case of DF, the stem_c _1.30 relationship and stand density were affected by changes in soil fertility. The general fertilisation effect was an increment of 40% of ligneous dry matter for DF and only 22% for NS. In both cases, the amount of wood biomass produced per unit of leaf biomass (on a tree basis and, to a lesser extent, on a per hectare basis) was greater in fertilised plots. However, in the case of NS, the same amount of wood biomass was produced from a smaller quantity of leaves while in the case of DF, the same amount of leaves produced more wood biomass.The amount of nutrients in total ligneous biomass was higher for N, P and K, but lower for Ca and Mg, in DF than in NS. A high variability was observed between nutrient content of the different compartments, e.g., DF < NS for needles (except Mg), DF < NS for K, Ca and Mg for stem wood and DF > NS for N and P of stem wood. Fertilisation did not considerably change the hierarchy. On the basis of this study, all the indexes concerning stand production, wood density, nutrient use efficiency and response to fertilisation gave a net advantage to DF. This information is highly relevant for both ecological and practical purposes.     

Effect of wood ash application on the biomass distribution and physiological state of Norway spruce seedlings on sandy soils

The effect of wood ash applied as a fertiliser (0.25, 0.5 and 1 kg m^–2) to nutrient poor sandy soil on 4-year-old Norway spruce (Picea abies) growing in small-scale sample plots was studied. Analyses carried out with roots, stems, needles and shoots of different age showed that an increase in the pH level and K and Ca concentrations and a decrease in N and P concentrations in the soil was accompanied by a disbalance of nutrients in tree compartments. Stimulation of pigment synthesis in Norway spruce needles was observed, and no disbalances occurred in the pigment system (Chl a/Chl b, TChl/Car) guaranteeing normal process of photosynthesis. Biomass responses to wood ash application depended on the age of needles and shoots, being only slightly notable in current year organs. Inhibition of height growth of seedlings, but stimulation of root biomass was established.     

生境变化对栓皮栎幼苗营养元素含量的影响

以栓皮栎在我国天然分布的南界、中部和北界为研究样点,将不同纬度的栓皮栎实生幼苗移栽到同一地点,于生长旺季同时在种源地和移栽地进行取样和测定,探讨生境变化对栓皮栎幼苗生长旺季营养元素含量特征的影响.结果表明:生境变化对栓皮栎幼苗各器官N含量和茎P含量的影响显著,而对各器官K含量和叶片、根P含量影响较小;在种源地,栓皮栎各器官的N含量与纬度间呈极显著的正相关,茎、根的全P含量随纬度升高而降低,叶片N/P的变化不明显;移栽后,北界栓皮栎幼苗各器官的N、P含量显著降低,不同纬度来源对栓皮栎幼苗N、P含量的影响更为明显;各纬度栓皮栎叶片的N/P均有不同程度的升高.不同纬度栓皮栎幼苗不同器官的营养元素含量存在差异,其对生境变化的响应也不相同.     

Colony size, cell number, carbon and nitrogen contents of Phaeocystis pouchetii from western Norway

Phaeocystis pouchetii is an ecologically important colony-formingmarine phytoplankton species in northern hemisphere cold waters.It plays a central role in the processing of biogeochemicallyimportant elements in the upper ocean during spring blooms.Here, we report highly significant quantitative relationshipsamong colony size, cell number and particulate carbon and nitrogencontents of field populations of P. pouchetii, which providethe means to quantitatively convert its biological expressioninto units of biogeochemical significance. Populations weresampled from mesocosms incubated in situ in western Norway andeither fertilized with nitrate and phosphate or left unamended.Phaeocystis colony blooms developed in both scenarios, but weremuch greater in fertilized treatments. Colonies from the lattertreatments were larger, contained higher concentrations of cellsand more particulate carbon and nitrogen than those in the unfertilizedmesocosm. Considering all data, log cell number increased linearlywith log colony volume with a slope of 0.54. Log carbon andnitrogen increased with log colony volume, with respective slopesof 0.92 and 1.22, indicating a significant component of non-cellularcarbon and nitrogen within the colonies. Carbon and nitrogencontents of colonies were linearly related, and fertilized coloniescontained more nitrogen relative to carbon than unfertilizedcolonies. These equations are particularly applicable to contemporarystudies of P. pouchetii because they represent colonies growingin environments that mimic a continuum from natural to eutrophicatedecosystems.     

Effects of wood ash and nitrogen fertilization on fine root biomass and soil and foliage nutrients in a Norway spruce stand in Finland

We determined the effects of wood ash fertilization, given together with nitrogen (WAN), and nitrogen given together with P, B and Cu (SSF), on soil and foliage nutrients and fine root biomass in a 45-year-old Norway spruce stand in southern Finland. Fine roots were sampled 9 years, and the soil 10 years after ash (3 t/ha) and nitrogen (150 kg/ha) application. Fine root biomass tended to be lower, the necromass higher, and the fine root distribution relatively deeper on the WAN than on the control and SSF plots. The response of fine root biomass to WAN was probably related to changes in soil acidity. pH, base saturation, total and extractable concentrations of Ca, K, Mg and P, and total B, Cd, Mn, Ni and Zn concentrations in the organic layer were significantly higher on the WAN plots than on the SSF and the control plots with no ash and nutrient addition. On the WAN plots, the pH was 1.2 pH-units higher, the exchangeable Ca concentrations fourfold and those of Mg over twofold compared to the control plots. WAN increased the concentrations of K but decreased those of Mn and Ni in the needles compared to the control and SSF treatment. Even though ash and nitrogen fertilisation tended to decrease the fine root biomass, this decrease was not likely to affect tree growth during a 10-year period.     

Effects of long-term open-air exposure to fluoride,nitrogen compounds and SO2 on visible symptoms,pollutant accumulation and ultrastructure of Scots pine and Norway spruce seedlings

 Effects of SO₂, aqueous fluoride (NaF) and a solution of nitrogen compounds (NH₄NO₃) on the visible symptoms, pollutant accumulation and ultrastructure of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] seedlings were studied in an open-air experiment lasting for 3 consecutive years. Visible injury symptoms were most pronounced in combination exposures and whenever F was applied. Visible symptoms correlated well with needle pollutant concentrations. Exposure to NaF increased needle F contents particularly when F was applied with SO₂ or NH₄NO₃. This suggests that a reduction in N or SO₂ emissions, in F polluted areas, could improve the condition of conifers via decreased accumulation of phytotoxic F in the needles. Norway spruce needles accumulated 2 – 10 times as much S and F as those of Scots pine. Microscopic observations showed various changes in the needle mesophyll cell ultrastructure. In both species, exposure to SO₂ increased significantly the amount of cytoplasmic vacuoles, suggesting detoxification of excess sulphate or low pH. F treatments resulted in a significant enlargement of plastoglobuli in Scots pine and a darkening of plastoglobuli in Norway spruce. All exposures enhanced the accumulation of lipid bodies. An increased portion of translucent plastoglobuli was most pronounced in N treatments. Many of the ultrastructural changes and visible symptoms appeared only as number of years exposed increased, indicating that long-term experiments are needed. Both visible symptoms and ultrastructural changes pointed to the more pronounced sensitivity of Norway spruce compared to Scots pine. Ultrastructural results mostly supported earlier qualitative observations of F, N and SO₂ effects on needle mesophyll cell ultrastructure. However, no reduction of thylakoids in SO₂ containing exposure or curling of thylakoids in F exposure could be detected in the present study. Received: 5 December 1994 / Accepted: 28 April 1995     

The architecture of Norway spruce ectomycorrhizae: three-dimensional models of cortical cells, fungal biomass, and interface for potential nutrient exchange

Gathering realistic data on actual fungal biomass in ectomycorrhized fine root systems is still a matter of concern. Thus far, observations on architecture of ectomycorrhizae (ECMs) have been limited to analyses of two-dimensional (2-D) images of tissue sections. This unavoidably causes stereometrical problems that lead to inadequate assumptions about actual size of cells and their arrangement within ECM's functional compartments. Based on extensive morphological investigations of field samples, we modeled the architectural components of an average-sized Norway spruce ECM. In addition to our comprehensive and detailed quantitative data on cell sizes, we studied actual shape and size, in vivo arrangement, and potential nutrient exchange area of plant cortical cells (CCs) using computer-aided three-dimensional (3-D) reconstructions based on semithin serial sections. We extrapolated a factual fungal biomass in ECMs (Hartig net (HN) included) of 1.71 t?ha^?1 FW (0.36 t?ha^?1 DW) for the top 5 cm of soil for an autochthonous, montane, optimum Norway spruce stand in the Tyrolean Alps. The corresponding potential nutrient exchange area in ECMs including main axes of ECM systems, which is defined as the sum of interfaces between plant CCs and the HN, amounts to at least 3.2?×?10⁵?m²?ha^?1. This is the first study that determines the contribution of the HN to the total fungal biomass in ECMs as well as the quantification of its contact area. Our results may stimulate future research on fungal below-ground processes and their impact on the global carbon cycle.     

Endogenous levels of oxygen,carbon dioxide and ethylene in stems of Norway spruce trees during one growing season

Summary The trees sampled in this study came from two stands of Norway spruce, Picea abies (L.) Karst., near Stockholm, Sweden, differing in mean age and height. Holes were bored perpendicular to the stem surface, and gas samples were taken from the outer part of the sapwood throughout one growing season. Endogenous levels of molecular oxygen (O₂), carbon dioxide (CO₂) and ethylene in the outer sapwood were determined by combined gas chromatography — mass spectrometry (GC-MS) and GC. O₂ concentrations began to decrease as growth started in spring. The lowest levels (<5%) were recorded around mid-summer. In the younger stand concentrations remained below 5% until September. In October, O₂ concentrations in the sapwood were similar to those of air. Concentrations of CO₂ were below 1% in spring, but began to rise in May, peaking a month later at approximately 10%. Thereafter a slow decrease occurred until October, by which time levels had returned to those recorded in spring. Ethylene concentrations in the older stand reached 75 ppm early in May, while levels in the younger stand peaked at around 30 ppm later in May. Thereafter ethylene levels in both stands started to decrease down to ppb levels. The correlation between determined gas levels and physiological events associated with the seasonal growth cycle in temperate zones is discussed.     

热胁迫后施用氮肥和秸秆对土壤微生物生物量及功能的影响

研究了热胁迫(40℃,18 h)后在60 d培养期内不施物料(S)、仅施尿素(N)、仅施秸秆(R)和二者配施(RN)处理下土壤微生物生物量及土壤功能(基础呼吸、基质诱导呼吸、秸秆分解能力)的变化.结果表明:热胁迫有促进土壤微生物生物量及功能的趋势,但其影响弱且持续时间短.不论热胁迫与否,与不施物料和仅施氮肥处理相比,施用秸秆及其与氮肥配施处理下土壤微生物生物量、基础呼吸、基质诱导呼吸和秸秆分解能力均大幅提高,而单施氮肥处理与不施物料处理相比变化不明显,各项指标甚至有降低趋势.表明施用秸秆及其与氮肥配施对自然土壤和环境胁迫(干扰)后土壤功能都有改善作用.     

Effects of elevated CO2 and nitrogen on nutrient uptake in ponderosa pine seedlings

This paper reports on the results of a controlled-environment study on the effects of CO₂ (370, 525, and 700 mol mol^-1) and N [0, 200, and 400 g N g soil^-1 as (NH₄)SO₄] on ponderosa pine (Pinus ponderosa) seedlings. Based upon a review of the literature, we hypothesized that N limitations would not prevent a growth response to elevated CO₂. The hypothesis was not supported under conditions of extreme N deficiency (no fertilizer added to a very poor soil), but was supported when N limitations were less severe but still suboptimal (lower rate of fertilization). The growth increases in N-fertilized seedlings occurred mainly between 36 and 58 weeks without any additional N uptake. Thus, it appeared that elevated CO₂ allowed more efficient use of internal N reserves in the previously-fertilized seedlings, whereas internal N reserves in the unfertilized seedlings were insufficient to allow this response. Uptake rates of other nutrients were generally proportional to growth. Nitrogen treatment caused reductions in soil exchangeable K⁺, Ca²⁺, and Mg²⁺ (presumably because of nitrification and NO₃ ^- leaching) but increases in extractable P (presumably due to stimulation of phosphatase activity).The results of this and other seedling studies show that elevated CO₂ causes a reduction in tissue N concentration, even under N-rich conditions. The unique response of N is consistent with the hypothesis that the efficiency of Rubisco increases with elevated CO₂. These results collectively have significant implications for the response of mature, N-deficient forests to evevated CO₂.     

Effects of nitrogen supply and elevated carbon dioxide on construction cost in leaves of Pinus taeda (L.) seedlings

Seedlings of loblolly pine (Pinus taeda L.) were grown under varying conditions of soil nitrogen and atmospheric carbon dioxide availability to investigate the interactive effects of these resources on the energetic requirements for leaf growth. Increasing the ambient CO₂ partial pressure from 35 to 65 Pa increased seedling growth only when soil nitrogen was high. Biomass increased by 55% and photosynthesis increased by 13% after 100 days of CO₂ enrichment. Leaves from seedlings grown in high soil nitrogen were 7.0% more expensive on a g glucose g^–1 dry mass basis to produce than those grown in low nitrogen, while elevated CO₂ decreased leaf cost by 3.5%. Nitrogen and CO₂ availability had an interactive effect on leaf construction cost expressed on an area basis, reflecting source-sink interactions. When both resources were abundant, leaf construction cost on an area basis was relatively high (81.8±3.0 g glucose m^–2) compared to leaves from high nitrogen, low CO₂ seedlings (56.3±3.0 g glucose m^–2) and low nitrogen, low CO₂ seedlings (67.1±2.7 g glucose m^–2). Leaf construction cost appears to respond to alterations in the utilization of photoassimilates mediated by resource availability.