Elevated CO2 reduces field decomposition rates of Betula pendula (Roth.) leaf litter

The effect of elevated atmospheric CO₂ and nutrient supply on elemental composition and decomposition rates of tree leaf litter was studied using litters derived from birch (Betula pendula Roth.) plants grown under two levels of atmospheric CO₂ (ambient and ambient +250 ppm) and two nutrient regimes in solar domes. CO₂ and nutrient treatments affected the chemical composition of leaves, both independently and interactively. The elevated CO₂ and unfertilized soil regime significantly enhanced lignin/N and C/N ratios of birch leaves. Decomposition was studied using field litter-bags, and marked differences were observed in the decomposition rates of litters derived from the two treatments, with the highest weight remaining being associated with litter derived from the enhanced CO₂ and unfertilized regime. Highly significant correlations were shown between birch litter decomposition rates and lignin/N and C/N ratios. It can be concluded, from this study, that at levels of atmospheric CO₂ predicted for the middle of the next century a deterioration of litter quality will result in decreased decomposition rates, leading to reduction of nutrient mineralization and increased C storage in forest ecosystems. However, such conclusions are difficult to generalize, since tree responses to elevated CO₂ depend on soil nutritional status.     

Growth performance of hybrid families by crossing selfed lines of Betula pendula Roth

Hybrid breeding is an effective approach in many agricultural crops. In allogamous tree species severe inbreeding depression and long reproductive cycles generally prohibit its use. However, three generations of selfing in silver birch (Betula pendula Roth) were obtained by forcing trees to flowering under greenhouse conditions. Hybrids were produced by crossing first-, second and third-generation selfed lines. The effects of different levels of parental inbreeding on the growth performance of hybrid families were observed in a 9-year-old field progeny test. Also, provenance crosses were carried out between selfed lines from different parts of Finland and several other European countries. Observations of growth performance of the provenance hybrids were made in the same trial. The results indicated that the mean stem volumes were significantly different between classes of parental in breeding coefficients (F_P) (P<0.0001), and were positively correlated with F_P (r=0.9106, P<0.05). Within-family variation of the hybrid families decreased with an increase of F_P. The performance of the provenance crosses between parents at a relatively close distance did not depart significantly from the standard controls. However, when the cross distance was extended far to the south, hybrids grew faster, indicating either higher heterozygosity or an extended growth period.     

Cadmium and copper interactions on the accumulation and distribution of Cd and Cu in birch (Betula pendula Roth) seedlings

The effect of different external cadmium (Cd) and copper (Cu) regimes on the concentration of Cd and Cu in roots and shoots of birch (Betula pendula Roth.) seedlings was investigated. The seedlings were grown for 12 days in a weak nutrient solution (containing all essential nutrient elements including 0.025 µM Cu) at pH 4.2 with combinations of additional 0–2 µM CdCl₂ and 0–2 µM CuCl₂. Root and shoot concentrations of Cu were decreased by Cd in all treatments which included 0.1–2 µM of additional Cu in the treatment solution. When no extra Cu was added, only the shoot concentration of Cu was decreased by Cd whereas the root concentration was not affected. The shoot concentration of Cd was decreased by 0.5 and 2 µM of additional Cu in the treatment solution. The root concentration of Cd was decreased by Cu only when the concentration of additional Cu in the treatment solution was equal to or exceeded the concentration of Cd.     

Systematic within-tree variation in mountain birch leaf quality for a geometrid, Epirrita autumnata

Abstract.

• 1 We studied within-tree variation in leaf quality of the mountain birch, Betula pubescens ssp. tortuosa, for larvae of the autumnal moth, Epirrita autumnata.
• 2 The purpose of the study was to determine the possible occurrence of systematic differences in larval growth on short shoot leaves (i.e. leaves of the same age): among leaves facing in different compass directions, between leaves of lower and upper branches, among leaves on different positions within a branch and among leaves of different sizes within a short shoot. We also measured larval growth between short shoot and long shoot leaves (i.e. between leaves of different age).
• 3 The larvae grew best on leaves on the north side of trees and most poorly on south side leaves, the east and west sides being intermediate. Leaves from the upper branches supported larval growth better than leaves from the lower ones. The larvae grew better on the smallest leaf of each short shoot and were able to utilize it more efficiently than the two larger leaves. Short shoot leaves from the basal and middle parts of the upper branches of the trees were of better quality for the larvae than short shoot leaves from the tip part of the branches. The larval growth rate did not differ between short shoot and long shoot leaves. In general, within-tree variation in the larval growth rate was lower than variation among different trees.
• 4 Damage to leaves can decrease leaf quality for herbivores in the same year (rapidly inducible responses) or the following year(s) (delayed inducible responses). Our results show that systematic within-tree variation in larval growth can be as great as the effects of rapidly inducible responses and that variation among individual trees can be as great as the mean effects of delayed inducible responses.

     

Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling

The response of temperate forest ecosystems to elevated atmospheric CO₂ concentrations is important because these ecosystems represent a significant component of the global carbon cycle. Two important but not well understood processes which elevated CO₂ may substantially alter in these systems are regeneration and nitrogen cycling. If elevated CO₂ leads to changes in species composition in regenerating forest communities then the structure and function of these ecosystems may be affected. In most temperate forests, nitrogen appears to be a limiting nutrient. If elevated CO₂ leads to reductions in nitrogen cycling through increased sequestration of nitrogen in plant biomass or reductions in mineralization rates, long-term forest productivity may be constrained. To study these processes, we established mesocosms of regenerating forest communities in controlled environments maintained at either ambient (375 ppm) or elevated (700 ppm) CO₂ concentrations. Mesocosms were constructed from intact monoliths of organic forest soil. We maintained these mesocosms for 2 years without any external inputs of nitrogen and allowed the plants naturally present as seeds and rhizomes to regenerate. We used ¹⁵N pool dilution techniques to quantify nitrogen fluxes within the mesocosms at the end of the 2 years. Elevated atmospheric CO₂ concentration significantly affected a number of plant and soil processes in the experimental regenerating forest mesocosms. These changes included increases in total plant biomass production, plant C/N ratios, ectomycorrhizal colonization of tree fine roots, changes in tree fine root architecture, and decreases in plant NH₄ ⁺ uptake rates, gross NH₄ ⁺ mineralization rates, and gross NH₄ ⁺ consumption rates. In addition, there was a shift in the relative biomass contribution of the two dominant regenerating tree species; the proportion of total biomass contributed by white birch (Betula papyrifera) decreased and the proportion of total biomass contributed by yellow birch (B. alleghaniensis) increased. However, elevated CO₂ had no significant effect on the total amount of nitrogen in plant and soil microbial biomass. In this study we observed a suite of effects due to elevated CO₂, some of which could lead to increases in potential long term growth responses to elevated CO₂, other to decreases. The reduced plant NH₄ ⁺ uptake rates we observed are consistent with reduced NH₄ ⁺ availability due to reduced gross mineralization rates. Reduced NH₄ ⁺ mineralization rates are consistent with the increases in C/N ratios we observed for leaf and fine root material. Together, these data suggest the positive increases in plant root architectural parameters and mycorrhizal colonization may not be as important as the potential negative effects of reduced nitrogen availability through decreased decomposition rates in a future atmosphere with elevated CO₂. Received: 10 January 1997 / Accepted: 25 July 1997     

Carbon allocation and carbon transfer between t Betula papyrifera and t Pseudotsuga menziesii seedlings using a 13C pulse-labeling method

Here we describe a simple method for pulse-labeling tree seedlings with ¹³CO_2(gas), and then apply the method in two related experiments: t (i) comparison of carbon allocation patterns between t Betula papyrifera Marsh. and t Pseudotsuga menziesii (Mirb.) Franco, and t (ii) measurement of one-way belowground carbon transfer from t B. papyrifera to t P. menziesii. Intraspecific carbon allocation patterns and interspecific carbon transfer both influence resource allocation, and consequently development, in mixed communities of t B. papyrifera and t P. menziesii.In preparation for the two experiments, we first identified the appropriate ¹³CO_2(gas) pulse-chase regime for labeling seedlings: a range of pulse (100-mL and 200-mL 99 atom%¹³ CO_2(gas)) and chase (0, 3 and 6 d) treatments were applied to one year-old t B. papyrifera and t P. menziesii seedlings. The amount of ¹³CO₂ fixed immediately after 1.5 h exposure was greatest for both t B. papyrifera (40.8 mg excess ¹³C) and t P. menziesii (22.9 mg excess ¹³C) with the 200-mL pulse, but higher ¹³C loss and high sample variability resulted in little difference in excess¹³ C content between pulse treatments after 3 d for either species. The average excess ¹³C root/shoot ratio of t B. papyrifera and t P. menziesii changed from 0.00 immediately following the pulse to 0.61 and 0.87 three and six days later, which reflected translocation of 75% of fixed isotope out of foliage within 3 d following the pulse and continued enrichment in fine roots over 6 d. Based on these results, the 100-mL CO_2(gas) and 6-d chase were considered appropriate for the carbon allocation and belowground transfer experiments.In the carbon allocation experiment, we found after 6 d that t B. papyrifera allocated 49% (average 9.5 mg) and t P. menziesii 41% (average 5.8 mg) of fixed isotope to roots, of which over 55% occurred in fine roots in both species. Species differences in isotope allocation patterns paralleled differences in tissue biomass distribution. The greater pulse labeling efficiency of t B. papyrifera compared to t P. menziesii was associated with its two-fold and 13- fold greater leaf and whole seedling net photosynthetic rates, respectively, 53% greater biomass, and 35% greater root/shoot ratio.For the carbon transfer experiment, t B. papyrifera and t P. menziesii were grown together in laboratory rootboxes, with their roots intimately mingled. A pulse of 100 mL¹³ CO_2(gas) was applied to paper birch and one-way transfer to neighboring t P. menziesii was measured after 6 d. Of the excess ¹³C fixed by t B. papyrifera, 4.7% was transferred to neighboring t P. menziesii, which distributed the isotope evenly between roots and shoots. Of the isotope received by t P. menziesii, we estimated that 93% was taken up through belowground pathways, and the remaining 7% taken up by foliage as¹³ CO_2(gas) respired by t B. papyrifera shoots. These two experiments indicate that t B. papyrifera fixes more total carbon and allocates a greater proportion to its root system than does t P. menziesii, giving it a competitive edge in resource gathering; however, below-ground carbon sharing is of sufficient magnitude that it may help ensure co-existence of the two species in mixed communities.     

Phenolic compounds in seedlings of Betula pubescens and B. pendula are affected by enhanced UVB radiation and different nitrogen regimens during early ontogeny

We studied the ability of tree seedlings to respond to two environmental factors, elevated ultraviolet B (UVB) radiation and availability of nitrogen (N), at the beginning of their development. Seeds of two birch species, Betula pubescens Ehrh. (common white birch) and B. pendula Roth (silver birch), were germinated and the seedlings grown in an experimental field in eastern Finland. The experimental design consisted of a constant 50% increase in UVB radiation (including a slight increase in UVA), a UVA control (a slight increase in UVA) and a control. The seedlings were fertilized with three levels of N. The experiment lasted for 2 months; aboveground biomass was measured and the most mature leaf of each seedling was taken for the analyses of phenolics. Growth of the seedlings was not significantly affected by enhanced UVB, but was increased by increasing N. Elevated UVB induced significant changes in phenolic compounds. Quercetin glycosides were accumulated in the leaves of both species in response to UVB; this is considered to be a protective response. However, the direction of the responses of individual phenolics to different N regimens differed. In addition, concentration of soluble condensed tannins was lower at moderate N than that at lower levels of N in both species; on the contrary, in B. pubescens the concentration of insoluble condensed tannins was highest at moderate N. No significant interaction between UV and N was detected, and the responses of the two species were highly similar to UVB, while the responses to N regimens varied slightly more between species.     

Soil CO2 efflux of two silver birch clones exposed to elevated CO2 and O3 levels during three growing seasons

In the present open‐top chamber experiment, two silver birch clones (Betula pendula Roth, clone 4 and clone 80) were exposed to elevated levels of carbon dioxide (CO₂) and ozone (O₃), singly and in combination, and soil CO₂ efflux was measured 14 times during three consecutive growing seasons (1999–2001). In the beginning of the experiment, all experimental trees were 7 years old and during the experiment the trees were growing in sandy field soil and fertilized regularly. In general, elevated O₃ caused soil CO₂ efflux stimulation during most measurement days and this stimulation enhanced towards the end of the experiment. The overall soil respiration response to CO₂ was dependent on the genotype, as the soil CO₂ efflux below clone 80 trees was enhanced and below clone 4 trees was decreased under elevated CO₂ treatments. Like the O₃ impact, this clonal difference in soil respiration response to CO₂ increased as the experiment progressed. Although the O₃ impact did not differ significantly between clones, a significant time × clone × CO₂× O₃ interaction revealed that the O₃‐induced stimulation of soil respiration was counteracted by elevated CO₂ in clone 4 on most measurement days, whereas in clone 80, the effect of elevated CO₂ and O₃ in combination was almost constantly additive during the 3‐year experiment. Altogether, the root or above‐ground biomass results were only partly parallel with the observed soil CO₂ efflux responses. In conclusion, our data show that O₃ impacts may appear first in the below‐ground processes and that relatively long‐term O₃ exposure had a cumulative effect on soil CO₂ efflux. Although the soil respiration response to elevated CO₂ depended on the tree genotype as a result of which the O₃ stress response might vary considerably within a single tree species under elevated CO₂, the present experiment nonetheless indicates that O₃ stress is a significant factor affecting the carbon cycling in northern forest ecosystems.     

Carbon transfer associated with assimilation of organic nitrogen sources by silver birch (Betula pendula Roth.)

Summary Qualitative and quantitative aspects of heterotrophic carbon assimilation by mycorrhizal plants of birch (Betula pendula) were examined. Plants were grown aseptically from seed in the mycorrhizal condition with the fungus Hebeloma crustuliniforme and in the non-mycorrhizal condition, with protein as their sole exogenous nitrogen source. Yields and nitrogen contents were determined in some of the plants, while the roots of others were supplied with ¹⁴C-labelled protein and their shoots exposed for up to 72 h to different irradiance regimes. Only mycorrhizal plants utilised the organic nitrogen. Uptake of carbon associated with this utilisation and its translocation to the leaves was demonstrated directly by means of autoradiography. Amounts of activity transferred to shoots were greatest in low irradiance regimes. Calculation of net carbon gain from the heterotrophic source, based upon the assumption that breakdown products of protein are assimilated as amino-acids, indicates that over a 55-day growth period up to 9% of plant C may be derived from protein. The physiological and ecological significance of these findings are discussed.