Differences in tree and shrub growth responses to climate change in a boreal forest in China

Global climate change has led to rising temperatures and drought in boreal forests in Northeast China. In some areas, shrubs and trees coexist in high altitude and high latitude areas, and their differences with global warming may lead to significant changes in vegetation composition and distribution. Therefore, we compared the relationships between climate and growth for the most widely distributed dwarf shrub (Pinus pumila) and the two dominant tree species (Larix gmelinii and Pinus sylvestris var. mongolica) in boreal forests in the Daxing’an Mountains, China. A total of 340 tree-ring cores from 172 trees and 64 discs from shrubs were collected from four sites and compared the responses of shrub and tree growth to climate patterns using dendrochronological methods. The shrub and two tree species responded differently to interannual climate variance. The negative effect of growing season temperature was greater on growth of L.gmelinii and P. sylvestrisvar.mongolica than on P. pumila, and the promoting effect of winter and spring precipitation was greatest on P. pumila. Compared with the two tree species, P. pumila had a higher temperature threshold and grew over a shorter growing season. Our findings suggested that L. gmelinii and P. sylvestrisvar.mongolica are more susceptible to global warming than the shrubs that coexist with them. However, P.pumila should be studied from an individual perspective in the future due to the dwarf morphology of shrubs and their complex microenvironment.     

DOM对米槠次生林不同土层土壤微生物呼吸及其熵值的影响

可溶性有机质(Dissolved organic matter,DOM)作为土壤可溶性有机碳的重要来源,进入土壤之后通过改变土壤微生物数量和活性影响土壤矿化。DOM输入对土壤微生物呼吸和熵值的研究多集中在表层土壤,但对深层土壤微生物呼吸和熵值的影响关注较少。通过室内培养实验(120 d)研究米槠(Castanopsis carlesii)鲜叶DOM添加对表层土壤(0—10 cm)和深层土壤(40—60 cm)微生物呼吸及其土壤代谢熵和微生物熵的影响,为揭示DOM输入对亚热带森林土壤碳过程的影响提供理论依据。结果表明,在培养第1天,添加DOM的表层和深层土壤CO_2瞬时排放速率均显著高于对照(P0.001),分别是对照(不添加DOM)的3.58倍和6.93倍,之后显著下降。就累积排放量而言,无论是DOM添加处理还是对照,表层土壤显著大于深层土壤;在米槠鲜叶DOM添加后,表层土壤累积排放量显著大于对照的表层土壤(P0.001),但DOM添加处理深层土壤累积排放量与对照的深层土壤无明显差异。就微生物生物量碳而言,表层土壤微生物生物量碳含量在培养期间显著大于深层土壤。在整个添加DOM培养期间,表层土壤微生物生物量碳含量显著大于表层对照土壤,深层土壤微生物生物量碳含量显著大于深层对照土壤(第3天除外)。培养结束时(120 d),米槠鲜叶DOM添加处理下,表层土壤和深层土壤有机碳含量与第3天相比分别减少26%和19%。米槠鲜叶DOM添加处理后的深层土壤代谢熵(qCO_2)显著低于对照的深层土壤和DOM添加处理的表层土壤qCO_2(P0.001),说明外源DOM进入深层土壤后提高了土壤微生物对碳的利用效率。米槠鲜叶DOM添加处理后的深层土壤微生物熵是培养第3天的1.58倍,显著大于培养初期(P0.05),而DOM添加处理的表层土壤、对照的表层土壤与深层土壤的微生物熵分别是培养第3天的68%、79%和21%,说明DOM添加提高了深层土壤质量。     

Effects of global climatic warming on the boreal forest

On the basis of the predictions of the global climatic warming induced by anthropogenic activities, as provided by climatologists, current state of knowledge regarding possible ecological consequences of the warming on the boreal biome was discussed. A 600 to 700 km northward advance of the biome along with the warming was predicted. Such a shift could take place for half a century or so, which would be an unprecedentedly fast rate of progression. This might cause a serious disorder in species composition of the biome, particularly in the boundary regions. As to the carbon sink or source issues, considerable uncertainties and knowledge gaps existed. Elevated temperature and CO₂ levels would stimulate photosynthesis to result in an increase of CO₂ uptake, while the temperature increase would promote decomposition of organic matter especially that stored in the soils to release CO₂ to the atmosphere. Behaviors of northern peat bogs, whereca. 700 Gt of organic matter was thought to be accumulated, would seriously affect the balance. However, overall ecosystematic carbon balance was yet to be fully studied. It was realized that multifunctional approaches needed to be developed so as to integrate pieces of various information into a holistic picture. Need for international collaboration research efforts was also addressed.     

Decomposition of soil organic matter from boreal black spruce forest: environmental and chemical controls

Black spruce forests are a dominant covertype in the boreal forest region, and they inhabit landscapes that span a wide range of hydrologic and thermal conditions. These forests often have large stores of soil organic carbon. Recent increases in temperature at northern latitudes may be stimulating decomposition rates of this soil carbon. It is unclear, however, how changes in environmental conditions influence decomposition in these systems, and if substrate controls of decomposition vary with hydrologic and thermal regime. We addressed these issues by investigating the effects of temperature, moisture, and organic matter chemical characteristics on decomposition of fibric soil horizons from three black spruce forest sites. The sites varied in drainage and permafrost, and included a “Well Drained” site where permafrost was absent, and “Moderately well Drained” and “Poorly Drained” sites where permafrost was present at about 0.5 m depth. Samples collected from each site were incubated at five different moisture contents (2, 25, 50, 75, and 100% saturation) and two different temperatures (10°C and 20°C) in a full factorial design for two months. Organic matter chemistry was analyzed using pyrolysis gas chromatography-mass spectrometry prior to incubation, and after incubation on soils held at 20°C, 50% saturation. Mean cumulative mineralization, normalized to initial carbon content, ranged from 0.2% to 4.7%, and was dependent on temperature, moisture, and site. The effect of temperature on mineralization was significantly influenced by moisture content, as mineralization was greatest at 20°C and 50–75% saturation. While the relative effects of temperature and moisture were similar for all soils, mineralization rates were significantly greater for samples from the “Well Drained” site compared to the other sites. Variations in the relative abundances of polysaccharide-derivatives and compounds of undetermined source (such as toluene, phenol, 4-methyl phenol, and several unidentifiable compounds) could account for approximately 44% of the variation in mineralization across all sites under ideal temperature and moisture conditions. Based on our results, changes in temperature and moisture likely have similar, additive effects on in situ soil organic matter (SOM) decomposition across a wide range of black spruce forest systems, while variations in SOM chemistry can lead to significant differences in decomposition rates within and among forest sites.     

Nonlinear responses of ecosystem carbon fluxes to nitrogen deposition in an old-growth boreal forest

Nitrogen (N) deposition is known to increase carbon (C) sequestration in N-limited boreal forests. However, the long-term effects of N deposition on ecosystem carbon fluxes have been rarely investigated in old-growth boreal forests. Here we show that decade-long experimental N additions significantly stimulated net primary production (NPP) but the effect decreased with increasing N loads. The effect on soil heterotrophic respiration (Rh) shifted from a stimulation at low-level N additions to an inhibition at higher levels of N additions. Consequently, low-level N additions resulted in a neutral effect on net ecosystem productivity (NEP), due to a comparable stimulating effect on NPP and Rh, while NEP was increased by high-level N additions. Moreover, we found nonlinear temporal responses of NPP, Rh and NEP to low-level N additions. Our findings imply that actual N deposition in boreal forests likely exerts a minor contribution to their soil C storage.     

A substrate-induced growth-response method for estimating the biomass of microbial functional groups in soil and aquatic systems

Abstract A substrate-induced growth-response (SIGR) method is presented for estimating the biomass or density of microorganisms capable of carrying out specific metabolic functions in natural and human-made systems. The biomass of active organisms can be estimated based on the concentration of substrate needed to induce the growth of the standing population. Curves of substrate mineralization or depletion are used as indirect indicators of growth. Estimates of population size are obtained by using non-linear regression techniques to fit simple models, that contain biologically relevant parameters, to the substrate mineralization curves. In the present study, the SIGR method was used to estimate the numbers of organisms capable of mineralizing 2,4-dinitrophenol in soil and in a model waste-treatment system. The SIGR approach was tested by comparison of model estimates to estimates for the same parameters obtained by independent means.     

Chemical composition of aquatic dissolved organic matter in five boreal forest catchments sampled in spring and fall seasons

The chemical composition and carbon isotope signature of aquatic dissolved organic matter (DOM) in five boreal forest catchments in Scandinavia were investigated. The DOM was isolated during spring and fall seasons using a reverse osmosis technique. The DOM samples were analyzed by elemental analysis, FT-IR, solid-state CP-MAS ¹³C-NMR, and C-1s NEXAFS spectroscopy. In addition, the relative abundance of carbon isotopes (¹²C, ¹³C, ¹⁴C) in the samples was measured. There were no significant differences in the chemical composition or carbon isotope signature of the DOM sampled in spring and fall seasons. Also, differences in DOM composition between the five catchments were minor. Compared to reference peat fulvic and humic acids, all DOM samples were richer in O-alkyl carbon and contained less aromatic and phenolic carbon, as shown by FT-IR, ¹³C-NMR, and C-1s NEXAFS spectroscopy. The DOM was clearly enriched in ¹⁴C relative to the NBS oxalic acid standard of 1950, indicating that the aquatic DOM contained considerable amounts of organic carbon younger than about 50 years. The weight-based C:N ratios of 31 ± 6 and the values of indicate that the isolated DOM is of terrestrial rather than aquatic origin. We conclude that young, hydrophilic carbon compounds of terrestrial origin are predominant in the samples investigated, and that the composition of the aquatic DOM in the studied boreal forest catchments is rather stable during low to intermediate flow conditions.     

Impact of forest conversion to agriculture on carbon and nitrogen mineralization in subarctic Alaska

Land-use change is likely to be a major component of global change at high latitudes, potentially causing significant alterations in soil C and N cycling. We addressed the biogeochemical impacts of land-use change in fully replicated black spruce forests and agricultural fields of different ages (following deforestation) and under different management regimes in interior Alaska. Change from forests to cultivated fields increased summer temperatures in surface soil layers by 4–5 °C, and lengthened the season of biological activity by two to three weeks. Decomposition of a common substrate (oat stubble) was enhanced by 25% in fields compared to forests after litter bags were buried for one year. In-situ net N mineralization rates in site-specific soil were similar in forests and fields during summer, but during winter, forests were the only sites where net N immobilization occurred. Field age and management had a significant impact on C and N mineralization. Rates of annual decomposition, soil respiration and summer net N mineralization tended to be lower in young than in old fields and higher in fallow than in planted young fields. To identify the major environmental factors controlling C and N mineralization, soil temperature, moisture and N availability were studied. Decomposition and net N mineralization seemed to be mainly driven by availability of inorganic N. Soil temperature played a role only when comparing forests and fields, but not in field-to-field differences. Results from soil respiration measurements in fields confirmed low sensitivity of heterotrophic respiration, and thus decomposition to temperature. In addition, both soil respiration and net N mineralization were limited by low soil water contents. Our study showed that (1) C and N mineralization are enhanced by forest clearing in subarctic soils, and (2) N availability is more important than soil temperature in controling C and N mineralization following forest clearing. Projecting the biogeochemical impacts of land-use change at high latitudes requires an improved understanding of its interactions with other factors of global change, such as changing climate and N deposition.     

Successional and physical controls on the retention of nitrogen in an undisturbed boreal forest ecosystem

Floristic succession in the boreal forest can have a dramatic influence on ecosystem nutrient cycling. We predicted that a decrease in plant and microbial demand for nitrogen (N) during the transition from mid- to late-succession forests would induce an increase in the leaching of dissolved inorganic nitrogen (DIN), relative to dissolved organic nitrogen (DON). To test this, we examined the chemistry of the soil solution collected from within and below the main rooting zones of mid- and late-succession forests, located along the Tanana River in interior Alaska. We also used a combination of hydrological and chemical analyses to investigate a key assumption of our methodology: that patterns of soil water movement did not change during this transition. Between stands, there was no difference in the proportion of DIN below the rooting zone. 84–98% of DIN at both depths consisted of nitrate, which was significantly higher in the deeper mineral soil than at the soil surface (0.46±0.12 mg NO⁻ ₃ –N l⁻¹ vs 0.17±0.12 mg NO⁻ ₃ –N l⁻¹, respectively), and 79–92% of the total dissolved N consisted of DON. Contrary to our original assumption that nutrients were primarily leached downward, out of the rooting zone, we found much evidence to suggest that the glacially-fed Tanana River (>200 m from these stands) was contributing to the influx of water and nutrients into the soil active layer of both stands. Soil water potentials were positively correlated with river discharge; and ionic and isotopic (δ¹⁸O of H₂O) values of the soil solution closely matched those of river water. Thus, our ability to elucidate biological control over ecosystem N retention was confounded by riverine nutrient inputs. Climatic warming is likely to extend the season of glacial melt and increase riverine nutrient inputs to forests along glacially-fed rivers.     

Mineral nitrogen and microbial dynamics in the forest floor of clearcut or partially harvested successional boreal forest stands

The effects of clearcut and partial harvesting of early-seral trembling aspen plots were compared to conventional clearcut harvesting in mid-seral mixedwood and late-seral conifer plots. Twice a year, for three consecutive years, we assessed mineral N and microbial dynamics in the forest floor of these plots to test three hypotheses related to the higher litter quality of aspen leaves and to the sustained inputs of available C on partially harvested plots: (1) the post-clearcutting mineral N flush and the net [(NO₃^–): (NO₃^– + NH₄⁺)] production ratio (RNI) are higher in aspen plots than in black spruce plots, with intermediate values occurring in mixedwood plots; (2) net N mineralization rates in aspen plots are higher in spring than in autumn; and (3) compared to clearcutting, partial harvesting reduces potential ammonification and nitrification rates. Initial NH₄⁺ and NO₃^– concentrations respectively ranged between 1.7–4.4 and 0.2–1.5 g N kg^–1 N_total, net ammonification and nitrification rates (30 d incubations) respectively ranged between 5.3–17.8 and 0.1–27.6 g N kg^–1 N_total, basal respiration ranged between 20.9–38.9 mg CO₂-C kg^–1 h^–1, and microbial biomass ranged between 6.1–8.7 g C_mic kg^–1. Although clearcutting increased NO₃^– concentrations in aspen plots, the balance of our results did not support our first hypothesis, because NH₄⁺ concentrations increased in conifer plots only, potential ammonification was unaffected by clearcutting, potential nitrification increased in mixedwood plots only, and RNI increased in all plots. In each seral stage, basal respiration, microbial biomass, and metabolic quotient either increased or were unaffected by clearcutting, suggesting that increases in RNI after disturbance were not related to lower microbial immobilisation of NO₃^– due to lower available C. Forest floors in mid-seral mixedwood plots exhibited a distinct combination of mineral N and microbial properties, suggesting that the functional richness of the forest is enhanced not only by the number of species, but also by the diversity of assemblages that are present. Results supported our second hypothesis and showed, furthermore, that net N mineralization in conifer stands is greater in autumn than in spring. Partial harvesting in aspen stands resulted in lower potential mineralization of N and lower RNI, compared to clearcutting. Further lysimetry studies are needed to confirm whether partial harvesting mitigates NO₃^– leaching following disturbance.     

Species-dependent responses of soil microbial properties to fresh leaf inputs in a subtropical forest soil in South China

Aims Forest disturbance from extreme weather events due to climate change could increase the contribution of fresh green leaves to the litter layer of soil and subsequently alter the composition and activity of the soil microbial properties and soil carbon cycling. The objective of this study was to compare the effect of naturally fallen litter and fresh leaves on the soil microbial community composition and their activities.Methods Fresh leaves and normal fallen litter were collected from four tree species (Pinus elliottii, Schima superba, Acacia mangium, A. auriculaeformis) in subtropical China and mixed with soil. Soil microbial community composition was determined using PLFAs, and its activity was quantified by soil respiration. During a 12-month period, the decomposition rate of litter was measured bimonthly using a litterbag method. Soil microbial samples were collected after 6 and 12 months. Soil respiration was measured monthly.Important findings We found that fresh leaves decomposed faster than their conspecific fallen litter. Although total microbial biomass and bacterial biomass were similar among treatments, soil fungal biomass was higher in fresh leaf than fallen litter treatments, resulting in greater values of the Fungal phospholipid fatty acids (PLFAs)/Bacterial PLFAs ratio. Fungal PLFA values were greater for Schima superba than the other species. The effect of litter type on soil respiration was species-dependent. Specifically, fallen litter released 35% more CO₂ than fresh leaves of the conifer P. elliottii. The opposite pattern was observed in the broadleaf species whose fresh leaf treatments emitted 17%–32% more CO₂ than fallen litter. Given future predictions that global climate change will cause more disturbances to forests, these results indicate that conifer and broadleaf forests in subtropical China may respond differently to increased fresh litter inputs, with net soil microbial respiration decreasing in conifer forests and increasing in broadleaf forests.     

The origin of soil organic matter controls its composition and bioreactivity across a mesic boreal forest latitudinal gradient

Warmer climates have been associated with reduced bioreactivity of soil organic matter (SOM) typically attributed to increased diagenesis; the combined biological and physiochemical transformation of SOM. In addition, cross‐site studies have indicated that ecosystem regime shifts, associated with long‐term climate warming, can affect SOM properties through changes in vegetation and plant litter production thereby altering the composition of soil inputs. The relative importance of these two controls, diagenesis and inputs, on SOM properties as ecosystems experience climate warming, however, remains poorly understood. To address this issue we characterized the elemental, chemical (nuclear magnetic resonance spectroscopy and total hydrolysable amino acids analysis), and isotopic composition of plant litter and SOM across a well‐constrained mesic boreal forest latitudinal transect in Atlantic Canada. Results across forest sites within each of three climate regions indicated that (1) climate history and diagenesis affect distinct parameters of SOM chemistry, (2) increases in SOM bioreactivity with latitude were associated with elevated proportions of carbohydrates relative to plant waxes and lignin, and (3) despite the common forest type across regions, differences in SOM chemistry by climate region were associated with chemically distinct litter inputs and not different degrees of diagenesis. The observed climate effects on vascular plant litter chemistry, however, explained only part of the regional differences in SOM chemistry, most notably the higher protein content of SOM from warmer regions. Greater proportions of lignin and aliphatic compounds and smaller proportions of carbohydrates in warmer sites' soils were explained by the higher proportion of vascular plant relative to moss litter in the warmer relative to cooler forests. These results indicate that climate change induced decreases in the proportion of moss inputs not only impacts SOM chemistry but also increases the resistance of SOM to decomposition, thus significantly altering SOM cycling in these boreal forest soils.     

A comparative account of the microbiological characteristics of soils under natural forest,grassland and cropfield from Eastern India

Microbiological and physico-chemical characteristics of tropical forest, grassland and cropfield soils from India were investigated. The study revealed that the conversion of natural forest led to a reduction of soil organic C (26–36%), total N (26–35%), total P (33–44%), microfungal biomass (44–66%) and total microbial biomass C, N and P (25–60%) over a period of 30–50 years. Comparative analysis of microbial activity in terms of basal soil respiration revealed maximum activity in the forest and minimum in the cropfield soil. Analysis of microbial metabolic respiratory activity (qCO₂) indicated relatively greater respiratory loss of CO₂-C per unit microbial biomass in cropfield and grassland than in forest soil. Considering the importance of the microbial component in soil, we conclude that the conversion of the tropical forest to different land uses leads to the loss of biological stability of the soil.     

A phytosociological survey of the boreal forest (shape Vaccinio-Piceetea) in North America

A survey of syntaxa of vegetation of North American boreal forests (class Vaccinio-Piceetea) is presented. This phytosociological survey, carried out combining the Braun-Blanquet method with numerical syntaxonomical analyses (cluster and correspondence analysis), describes the associations of the North American boreal forests, which have several species, varieties or vicariant species in common with their Eurasian counterparts, and can be placed in the class Vaccinio-Piceetea. By means of tabular and multivariate analyses, 2084 North American relevés were compared with 3273 relevés from European, Japanese and Korean boreal forest, to describe and typify 4 orders, 10 alliances and 37 associations. Diagnostic tables, ordination, clustering, and climatic, edaphic and biogeographical data were used to show floristic affinities among these syntaxa and interpret their distribution areas. Syntaxa were briefly characterized by their floristic composition, physiognomy, succession, zonation, and biogeographical distribution.     

Nitrogen acquisition from inorganic and organic sources by boreal forest plants in the field

A wide range of recent studies have indicated that organic nitrogen may be of great importance to plant nitrogen (N) nutrition. Most of these studies have, however, been conducted in laboratory settings, excluding important factors for actual plant uptake, such as competition, mycorrhizal associations and soil interactions. In order to accurately evaluate the importance of different N compounds to plant N nutrition, field studies are crucial. In this study, we investigated short- as well as long-term plant nitrogen uptake by Deschampsia flexuosa, Picea abies and Vaccinium myrtillus from ¹⁵NO₃^–, ¹⁵NH₄⁺ and (U-¹³C, ¹⁵N) arginine, glycine or peptides. Root N uptake was analysed after 6 h and 64 days following injections. Our results show that all three species, irrespective of their type of associated mycorrhiza (arbuscular, ecto- or ericoid, respectively) rapidly acquired similar amounts of N from the entire range of added N sources. After 64 days, P. abies and V. myrtillus had acquired similar amounts of N from all N sources, while for D. flexuosa, the uptake from all N sources except ammonium was significantly lower than that from nitrate. Furthermore, soil analyses indicate that glycine was rapidly decarboxylated after injections, while other organic compounds exhibited slower turnover. In all, these results suggest that a wide range of N compounds may be of importance for the N nutrition of these boreal forest plants, and that the type of mycorrhiza may be of great importance for N scavenging, but less important to the N uptake capacity of plants.     

Understorey plant and soil responses to disturbance and increased nitrogen in boreal forests

Question: How do N fertilization and disturbance affect the understorey vegetation, microbial properties and soil nutrient concentration in boreal forests? Location: Kuusamo (66°22′N; 29°18′E) and Oulu (65°02′N; 25°47′E) in northern Finland. Methods: We conducted a fully factorial experiment with three factors: site (two levels), N fertilization (four levels) and disturbance (two levels). We measured treatment effects on understorey biomass, vegetation structure, and plant, soil and microbial N and C concentrations. Results: The understorey biomass was not affected by fertilization either in the control or in the disturbance treatment. Fertilization reduced the biomass of deciduous Vaccinium myrtillus. Disturbance had a negative effect on the biomass of V. myrtillus and evergreen Vaccinium vitis‐idaea and decreased the relative proportion of evergreen species. Fertilization and disturbance increased the biomass of grass Deschampsia flexuosa and the relative proportion of graminoids. The amount of NH₄⁺ increased in soil after fertilization, and microbial C decreased after disturbance. Conclusions: Our results suggest that the growth of slow‐growing Vaccinium species and soil microbes in boreal forests are not limited by N availability. However, significant changes in the proportion of dwarf shrubs to graminoids and a decrease in the biomass of V. myrtillus demonstrate the susceptibility of understorey vegetation to N enrichment. N enrichment and disturbance seem to have similar effects on understorey vegetation. Consequently, increasing N does not affect the rate or the direction of recovery after disturbance. Moreover, our study demonstrates the importance of understorey vegetation as a C source for soil microbes in boreal forests.     

Recovery of the ectomycorrhizal community after termination of long-term nitrogen fertilisation of a boreal Norway spruce forest

Ectomycorrhizal fungi (ECM) are a fundamental component of boreal forests promoting tree growth and participating in soil nutrient cycling. Increased nitrogen (N) input is known to largely influence ECM communities but their potential recovery is not well understood. Therefore, we studied the effects of long-term N-fertilisation on ECM communities, and their recovery after termination of N treatment. Fungal ITS sequencing data indicated that N-fertilisation (34 kg N ha⁻¹ y⁻¹) for 46 y decreased the relative abundance of ECM species in the fungal community and suppressed originally dominating medium-distance fringe exploration types adapted to N-limited conditions, while the ECM diversity remained unaffected. In other plots, 23 y after termination of fertilisation at 73 kg N ha⁻¹ y⁻¹ for 23 y, the relative abundance of ECM species shifted closer to, but did not reach, control levels. These observations indicate only slow recovery of ECM community, likely due to a high soil N retention capacity.