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Soil labile organic carbon (LOC) responds rapidly to environmental changes and plays an important role in carbon cycle. In this study, the seasonal fluctuations in LOC, the activities of carbon-cycle related enzymes, and the bacterial and fungal communities were analyzed for soils collected from two forests, namely Betula albosinensis (Ba) and Picea asperata Mast. (Pa), in the Qinling Mountains of China. Results revealed that the seasonal average contents of microbial biomass carbon (MBC), easily oxidized organic carbon (EOC), and dissolved organic carbon (DOC) of Pa forest soil were 13.5%, 30.0% and 15.7% less than those in Ba soil. The seasonal average enzyme activities of β-1,4-glucosidase (βG), and β-1,4-xylosidase (βX) of Ba forest soils were 30.0% and 32.3% higher than those of Pa soil while the enzyme activity of cellobiohydrolase (CBH) was 19.7% lower. Furthermore, the relative abundance of Acidobacteria was significantly higher in summer than in winter, whereas the relative abundance of Bacteroidetes was higher in winter. Regarding the fungal communities, the relative abundance of Basidiomycota was lowest in winter, whereas Ascomycota predominated in the same season. In addition, the soil LOC was significantly positively correlated with the CBH, βG and βX activities. Changes in LOC were significantly correlated with Acidobacteria, Bacteroidetes and Basidiomycota. We conclude that the seasonal fluctuations in forest soil LOC fractions relied on carbon cycle-associated enzymatic activities and microorganisms, which in turn were affected by climatic conditions.

Aims Competition, temperature and nutrient are the most important determinants of tree growth in the cold climate on the eastern Tibetan Plateau. Although many studies have reported their individual effects on tree growth, little is known about how the interactions of competition with fertilization and temperature affect root growth. We aim to test whether climate warming and fertilization promote competition and to explore the functional strategies of Picea asperata in response to the interactions of these factors. Methods We conducted a paired experiment including competition and non-competition treatments under elevated temperature (ET) and fertilization. We measured root traits, including the root tip number over the root surface (RTRS), the root branching events over the root surface (RBRS), the specific root length (SRL), the specific root area (SRA), the total fine root length and area (RL and RA), the root tips (RTs) and root branching (RB) events. These root traits are considered to be indicators of plant resource uptake capacity and root growth. The root biomass and the nutrient concentrations in the roots were also determined. Important Findings The results indicated that ET, fertilization and competition individually enhanced the nitrogen (N) and potassium (K) concentrations in fine roots, but they did not affect fine root biomass or root traits, including RL, RT, RA and RB. However, both temperature and fertilization, as well as their interaction, interacting with competition increased RL, RA, RT, RB and nutrient uptake. In addition, the SRL, SRA, RTRS and RBRS decreased under fertilization, the interaction between temperature and competition decreased SRL and SRA, while the other parameters were not affected by temperature or competition. These results indicate that P. asperata maintains a conservative nutrient strategy in response to competition, climate warming, fertilization and their interactions. Our results improve our understanding of the physiological and ecological adaptability of trees to global change.

Picea asperata and P. crassifolia have sympatric ranges and are closely related, but the differences between these species at the plastome level are unknown. To better understand the patterns of variation among Picea plastomes, the complete plastomes of P. asperata and P. crassifolia were sequenced. Then, the plastomes were compared with the complete plastomes of P. abies and P. morrisonicola, which are closely and distantly related to the focal species, respectively. We also used these sequences to construct phylogenetic trees to determine the relationships among and between the four species as well as additional taxa from Pinaceae and other gymnosperms. Analysis of our sequencing data allowed us to identify 438 single nucleotide polymorphism (SNPs) point mutation events, 95 indel events, four inversion events, and seven highly variable regions, including six gene spacer regions (psbJ-petA, trnT-psaM, trnS-trnD, trnL-rps4, psaC-ccsA, and rps7-trnL) and one gene (ycf1). The highly variable regions are appropriate targets for future use in the phylogenetic reconstructions of closely related, sympatric species of Picea as well as Pinaceae in general.

In this study, leaf litters of Betula albo-sinensis and 5 coniferous species were used as samples. The B. albo-sinensis leaf litter was buried in soil and termly treated with the water extracts of five types of coniferous leaf litter for a 6-month simulation decomposition experiment. The dynamics of mass loss and nutrients (C, N, P, and K) content of leaf litter and the soil enzymatic activities were measured to investigate the effects of plant secondary metabolites (PSM) from coniferous leaf litters on the decomposition processes of B. albo-sinensis leaf litter. The results indicated that the extracts of Pinus tabuliformis, Platycladus orientalis, P. armandii and Larix principis-rupprechtii leaf litters significantly inhibited the whole decomposition process and overall nutrients release of B. albo-sinensis leaf litter, while the extracts of Picea asperata leaf litter exhibited no significant influence. The general suppression of PSM on the soil sucrase, carboxymethyl cellulase and β-glucosidase activities might be the main reason leading to the inhibitory effects of the extracts of P. tabuliformis, P. orientalis, P. armandii and L. principis-rupprechtii leaf litter. The species causing inhibitory effects, especially L. principis-rupprechtii, was not recommended to be planted mixed with B. albo-sinensis, or their planting density should be lower in the mixed forests.

At an ecosystem level, stand age has a significant influence on carbon storage (CS). Dragon spruce (Picea asperata Mast.) situated along the upper reaches of the Bailongjiang River in northwest China were categorized into three age classes (29–32 years, Y1; 34–39 years, Y2; 40–46 years, Y3), and age-related differences in total carbon storage (TCS) of the forest ecosystem were investigated for the first time. Results showed that TCS for the Y1, Y2, and the Y3 age groups were 323.64, 240.66 and 174.60 Mg ha−1, respectively. The average TCS of the three age groups was 255.65 Mg C ha−1, with above-ground biomass, below-ground biomass, litter, and soil in the top 0.6 m contributing 15.0%, 3.7%, 12.1%, and 69.2%, respectively. CS in soil and TCS of the Y1 age group both significantly exceeded those of the Y3 age group (P

Background The terminal branch orders of plant root systems are increasingly known as an ephemeral module. This concept is crucial to recognize belowground processes. However, it is unknown if root modules still exist after they die? Methods The decomposition patterns of the first five root orders were observed for 3 years using a branch-order classification, a litter-bag method and sequential sampling in a common subalpine tree species (Picea asperata) of southwestern China. Results Two root modules were observed during the 3-year incubation. Among the first five branch orders, the first three order roots exhibited temporal patterns of mass loss, nutrients and stoichiometry distinct from their woody mother roots throughout the experimental period. This study, for the first time, reported the decomposition pattern of each individual root order and found a similar decomposition dynamic among ephemeral root branches in a forest tree species. Conclusions Results from this study suggest that root modules may also exist after death, while more data are needed for confirmation. The findings may further advance our understanding of architecture-associated functional heterogeneity in the fine-root system and also improve our ability to predict belowground processes.

According to the forest resources inventory data for different periods and the latest estimation parameters of forest carbon reserves in China, the carbon reserves and carbon density of forest biomass in the Tibet Autonomous Region from 1999 to 2019 were estimated using the IPCC international carbon reserves estimation model. The results showed that, during the past 20 years, the forest area, forest stock, and biomass carbon storage in Tibet have been steadily increasing, with an average annual increase of 1.85×104 hm2, 0.033×107 m3, and 0.22×107 t, respectively. Influenced by geographical conditions and the natural environment, the forest area and biomass carbon storage gradually increased from the northwest to the southeast, particularly in Linzhi and Changdu, where there are many primitive forests, which serve as important carbon sinks in Tibet. In terms of the composition of tree species, coniferous forests are dominant in Tibet, particularly those containing Abies fabri, Picea asperata, and Pinus densata, which comprise approximately 45% of the total forest area in Tibet. The ecological location of Tibet has resulted in the area being dominated by shelter forest, comprising 68.76% of the total area, 64.72% of the total forest stock, and 66.34% of the total biomass carbon reserves. The biomass carbon storage was observed to first increase and then decrease with increasing forest age, which is primarily caused by tree growth characteristics. In over-mature forests, trees’ photosynthesis decreases along with their accumulation of organic matter, and the trees can die. In addition, this study also observed that the proportion of mature and over-mature forest in Tibet is excessively large, which is not conducive to the sustainable development of forestry in the region. This problem should be addressed in future management and utilization activities.

Simple Summary During a survey of pine wilt disease in 2017, thousands of dead pitch pines (Pinus koraiensis) were discovered in Fengcheng, Liaoning Province, China. Samples were taken from several regions of the bole, and Bursaphelenchus xylophilus was collected from the dead pitch pine samples. Monochamus saltuarius is an indigenous insect in this area, but there is no experimental evidence that M. saltuarius can transport B. xylophilus in China. The experimental results showed that M. saltuarius was the new vector of pine wilt disease in China. In laboratory experiments, M. saltuarius fed and transmitted B. xylophilus not only on pines but also on other conifers. Taxus, Juniperus, Sabina and Podocarpus are new tree species upon which B. xylophilus can be fed and transmitted by M. saltuarius. Picea pungens, Picea asperata, Abies fabri, A. holophylla, Larix principis-rupprechtii, L. olgensis, Juniperus formosana, Sabina chinensis and Podocarpus macrophyllus are new species of coniferous trees known to be able to be infected by B. xylophilus in a laboratory setting. This study raises awareness to prevent the disease from expanding and provides a reference for the control of pine wilt disease. Abstract Pine wilt disease was first discovered in Dongtang town, Liaoning Province, China, in 2017. However, no record of Monochamus alteratus existed in Fengcheng, where M. saltuarius is an indigenous insect, and no experimental evidence has thus far indicated that M. saltuarius can transport the Bursaphelenchus xylophilus in China. In this study, we investigated whether M. saltuarius is a vector of B. xylophilus in China. On the sixth day after eclosion, beetles began to transmit nematodes into the twigs. The transmission period of nematodes is known to be able to last for 48 days after beetle emergence. In laboratory experiments, M. saltuarius fed and transmitted B. xylophilus not only on pines but also on other non-Pinus conifers. The non-Pinus conifers preferred by M. saltuarius for feeding are Picea pungens, Picea asperata, and Abies fabri. The experimental results show that M. saltuarius functions as a vector of B. xylophilus in northeast China.

Although the quantitative assessment of plant- and microbial-derived carbon (C) chemical composition in soil profiles has been initially explored, the vertical distribution pattern of these two C sources and their relative contribution in SOC formation based on the insights related to the rhizosphere are still lacking. We quantified the divergent accumulation of microbial-derived C (i.e., microbial residues), plant-derived C (i.e., lipids and lignin phenols) and SOC in the rhizosphere at various depths (0–10 cm, 10–20 cm and 20–30 cm) in the upper mineral soil and analyzed its control factors in an alpine coniferous forest (Picea asperata. Mast). We further revealed the relative contribution of plant- or microbial-derived C to rhizosphere SOC in the soil profile. The contents of microbial- and plant-derived C and SOC in the rhizosphere decreased with soil depth and were mainly regulated by root biomass and microbial biomass. Moreover, the contribution of microbial-derived C dominated by fungal residues to rhizosphere SOC at each soil depth (more than 62%) was much higher than that of plant-derived C (less than 6%), implying that the soil microbial C pump was intensely stimulated in the rhizosphere. These results indicated that microbial-derived C was the main contributor of rhizosphere SOC at various depths in the upper mineral soil. Our findings provide direct experimental evidence for assessing the relative contribution of microbial- or plant-derived C to SOC in the soil profile from the perspective of the rhizosphere.

About 70 % of the total land area in the world are affected by soil freeze and thaw (FT) cycles. Root is the first organ of plant to sense soil environment and it is unclear how it copes with the soil FT. Based on the different functions of firstorder pioneer and fibrous roots in woody plants, we hypothesize that pioneer and fibrous roots respond differently. The experiment was conducted in a growth chamber using Picea asperata seedlings. We designed the FT based on field observation data. The physiological responses in fibrous and pioneer roots were examined. Fibrous roots had higher root vitality and N content, whereas pioneer roots exhibited higher total nonstructural saccharide content. The accumulation of O2 - under FT treatment was similar in the two types of roots. Pioneer roots showed higher osmolyte (especially proline) content, whereas fibrous roots had higher peroxidase activity. The present study confirmed that fibrous roots have stronger metabolism ability, whereas pioneer roots are the key storage organs. FT in the temperature range from -5 to 5 °C are mild and do not cause serious injury to roots. Pioneer roots have higher tolerance to soil FT in spring than fibrous roots. The roots have different strategies to FT: fibrous roots increase the antioxidant system, whereas pioneer roots accumulate more osmolytes. Such knowledge can help us to understand how roots of woody plants cope with soil FT.

Litter chemical quality regulates the distinct composition of the main microbial groups and ecoenzymatic stoichiometry. Microbes in spruce ( Picea asperata Mast.) and fir ( Abies faxoniana Rehd.) rather than birch ( Betula platyphylla Suk.) and rhododendron ( Rhododendron lapponicum (L.) Wahl.) can more easily adjust their physiological metabolism to acclimate to low N resources. Litter decomposition is the main pathway of nutrient cycling that bridges aboveground and underground material circulation and energy flow. Microorganisms are essential for the regulation of organic carbon decomposition and nutrient cycling. We sought to reveal whether litter chemical quality predominates forest floor microbial structure and function in different species and how their characteristics vary with litter decomposition stages. We measured litter substrate quality, microbial community structure, microbial biomass carbon (MBC) and nitrogen (MBN), extracellular enzyme activities and stoichiometric homeostasis of fresh litter (L), and fermentative (F) and humus (H) layers for these tree species. Overall, the enzyme activities and microbial biomass of birch and rhododendron were greater than those of spruce and fir. The microbial abundances of birch and rhododendron decreased with decomposition. Forest floor microbial nutrient limitation is generally restricted by N in subalpine forests, and ecoenzymatic stoichiometry is affected mainly by dissolved C/N/P stoichiometry. Stronger microbial C:N homeostasis (H′) was observed for spruce (5.56) and fir (4.17) than that for birch (1.82) and rhododendron (1.33). We conclude that litter chemical quality led to the disparity in forest floor microbial groups and ecoenzymatic stoichiometry for different tree species.

Forests are important in the global carbon cycle and it is necessary to quickly and accurately measure forest volume to estimate forest aboveground biomass (AGB) and aboveground carbon storage (AGC). In this paper, we used data from the eighth forest resources inventory of China to establish two stand volume models based on stand density and forest basal area for 37 arbor forest types (dominant species), and performed a comparative analysis to obtain the best model. Then the AGB, AGB density, AGC, and AGC density of the different forest types and regions were estimated by conversion function methods. The results showed that: (1) The volume model of tree height and forest basal area could better fit the natural growth process of forests, and 36 of the 37 forest types had R2 greater than 0.8, (2) The average AGB density of arbor forest in China was 95.03 Mg ha&minus, 1 and the average AGC density was 48.15 Mg ha&minus, 1 (3) Among forest types, Picea asperata Mast., Quercus spp., and Populus spp. had the highest AGB and AGC, while Cinnamomum camphora (L.) Presl, Pinus taiwanensis Hayata, and Pinus densiflora Sieb. et Zucc. had the lowest. The AGB density and AGC density of Phoebe zhennan S. Lee et F. N. Wei and Pinus densata Mast. were the highest, while those of Pinus densiflora Sieb. et Zucc., Pinus elliottii Engelmann, and Eucalyptus robusta Smith were the lowest. (4) Among regions, AGB and AGC ranging from high to low, were as follows: northwest, southwest, northeast, central south, east, and north. The northwest and southwest regions accounted for more than 70% of the country&rsquo, s AGB and AGC. The average AGB density and AGC density among the regions were 91.34 Mg ha&minus, 1 and 46.4 Mg ha&minus, 1, respectively. Ranging from high to low as follows: southwest, northwest, northeast, east, central south, and north. The methods used in this paper provide a basis for fast and accurate estimation of stand volume, and the estimates of AGB and AGC have important reference value for explaining the role of ecosystems in coping with global climate change in China.

Phenylalanine ammonia-lyase (PAL) is the crucial enzyme of the phenylpropanoid pathway, which plays an important role in plant disease resistance. To understand the function of PAL in Picea asperata, in this study, the full-length cDNA sequence of the PAL gene from this species was isolated and named PaPAL. The gene contains a 2160-bp open reading frame (ORF) encoding 720 amino acids with a calculated molecular weight of 78.7 kDa and a theoretical isoelectric point of 5.88. The deduced PaPAL protein possesses the specific signature motif (GTITASGDLVPLSYIA) of phenylalanine ammonia-lyases. Multiple alignment analysis revealed that PaPAL has high identity with other plant PALs. The tertiary structure of PaPAL was predicted using PcPAL from Petroselinum crispum as a template, and the results suggested that PaPAL may have a similar function to that of PcPAL. Furthermore, phylogenetic analysis indicated that PaPAL has a close relationship with other PALs from the Pinaceae species. The optimal expression condition of recombinant PaPAL in Escherichia coli BL21 (DE3) was 0.2 mM IPTG (isopropyl &beta, D-thiogalactoside) at 16 &deg, C for 4 h, and the molecular weight of recombinant PaPAL was found to be approximately 82 kDa. Recombinant PaPAL was purified and exhibited high PAL activity at optimal conditions of pH 8.6 and 60 &deg, C. Quantitative real-time PCR (qRT-PCR) showed the PaPAL gene to be expressed in all tissues of P. asperata tested, with the highest expression level in the needles. The PaPAL gene was induced by the pathogen (Lophodermium piceae), which caused needle cast disease, indicating that it might be involved in defense against needle cast disease. These results provide a basis for understanding the molecular mechanisms of the PAL gene in the process of P. asperata disease resistance.

Picea belongs to the Pinaceae family and is a famous commercial tree species because of its straight trunk and excellent timber traits. Recently, omics have been widely used for fundamental and mechanism studies on Picea plants. To improve the accessibility to omics and phenotypic data and facilitate further studies, we compiled the sequences of 2 chloroplast genomes (Picea crassifolia and Picea asperata) and 32 complete omics data sets, including 20 transcriptomes, 4 proteomes, 2 degradomes and 6 microRNAs from P. crassifolia, P. asperata, Picea balfouriana and Picea abies tissues under different treatments, in PICEAdatabase. In addition, phenotypic data on plant growth and wood property traits were collected from two field trials of P. crassifolia. PICEAdatabase also includes useful analysis tools, such as BLAST, DESeq2 and JBrowse, to assist with analyses.

Understanding the variations in soil and plants with stand aging is important for improving management measures to promote the sustainable development of plantations. However, few studies have been conducted on the dynamics of stoichiometric traits and carbon (C), nitrogen (N), and phosphorus (P) pools across Picea asperata Mast plantations of different ages in subalpine regions. In the present study, we examined the stoichiometric traits and C, N, and P stocks in different components of three different aged (22-, 32-, and 42-year-old) P. asperata plantations by plot-level inventories. We hypothesized that the stoichiometric traits in mineral soil could shape the corresponding stoichiometric traits in soil microbes, tree roots and foliage, and the C, N, and P stocks of the total P. asperata plantation ecosystem would increase with increasing stand age. Our results show that the N:P ratio in mineral soil was significantly correlated with that in tree foliage and herbs. Additionally, the C:N ratio and C:P ratio in mineral soil only correlated with the corresponding stoichiometric traits in soil microbes and forest floor, respectively. Both the fractions of microbial biomass C in soil organic C and microbial biomass N in soil total N decreased with increasing stand age. The C, N, and P stocks of the total ecosystem did not continuously increase across stand development. In particular, the P stock of the total ecosystem exhibited a trend of increasing first and then decreasing. The aboveground tree biomass C accounted for more than 55% of the total ecosystem C stock regardless of stand age. In contrast, mineral soil and forest floor were the major contributors to the total ecosystem N and P stocks in all stands. This study suggested that all three different stands were N limited, and the stoichiometric homeostasis in the roots of P. asperata was more stable than that in the foliage. In addition, the soil microbial community assembly may change with increasing stand age for P. asperata plantations in the subalpine region.

Summary Partial desiccation treatment (PDT) stimulates germination and enhances the conversion of conifer somatic embryos. To better understand the mechanisms underlying the responses of somatic embryos to PDT, we used proteomic and physiological analyses to investigate these responses during PDT in Picea asperata. Comparative proteomic analysis revealed that, during PDT, stress‐related proteins were mainly involved in osmosis, endogenous hormones, antioxidative proteins, molecular chaperones and defence‐related proteins. Compared with those in cotyledonary embryos before PDT, these stress‐related proteins remained at high levels on days 7 (D7) and 14 (D14) of PDT. The proteins that differentially accumulated in the somatic embryos on D7 were mapped to stress and/or stimuli. They may also be involved in the glyoxylate cycle and the chitin metabolic process. The most significant difference in the differentially accumulated proteins occurred in the metabolic pathways of photosynthesis on D14. Furthermore, in accordance with the changes in stress‐related proteins, analyses of changes in water content, abscisic acid, indoleacetic acid and H2O2 levels in the embryos indicated that PDT is involved in water‐deficit tolerance and affects endogenous hormones. Our results provide insight into the mechanisms responsible for the transition from morphologically mature to physiologically mature somatic embryos during the PDT process in P. asperata.

Aims Plant roots play a critical role in the uptake of nutrients, and nitrogen (N) absorption is considered as the first step and a pivotal process in N metabolism of plants. Our objective was to better understand the absorption of two major inorganic N forms (NH4 and NO3) in subalpine coniferous forests under global warming. Methods Experimental warming using infrared heater was applied to two dominant species in subalpine coniferous forests of Sichuan, China, Picea asperata and Abies fargesii var. faxoniana. The non-invasive micromeasurement technology was used to investigate the effects of warming on the uptake rates of NH4 and NO3 and the potential interactions between these two ions. Important findings Results showed that the maximal net root uptake of NH4 and NO3 occurred at a distance of 17–18 mm and 17 mm from root tips, respectively for P. asperata. and at a distance of 11 mm and 11.5 mm from root tips respectively for A. fargesii var. faxoniana. Experimental warming elevated the uptake rates of NH4 and NO3 in both species, but the interactions between NH4 and NO3 differed between the two species. While NO3 uptake was inhibited in the presence of NH4 for both P. asperata and A. fargesii var. faxoniana, net NH4 uptake was promoted by NO3 supply only in P. asperata roots under experimental warming.

Fine roots play a crucial role in plant survival potential and biogeochemical cycles of forest ecosystems. Subalpine areas of the Eastern Qinghai-Tibetan Plateau have experienced different forest re-establishment methods after clear-cutting primary forest. However, little is known about fine root dynamics of these forests originating from artificial, natural and their combined processes. Here, we determined fine root traits (biomass, production and turnover rate) of three subalpine forest types, i.e., Picea asperata Mast. plantation forest (artificial planting, PF), natural secondary forest (natural without assisted regeneration, NF) and P. asperata broadleaved mixed forest (natural regeneration after artificial planting, MF) composed of planted P. asperata and naturally regenerated native broadleaved species. At the soil depth of 0&ndash, 30 cm, fine root biomass was the highest in PF and fine root production was the highest in NF, and both were the lowest in MF. Fine root dynamics of the three forest types tended to decrease with soil depth, with larger variations in PF. Fine root biomass and production were the highest in PF in 0&ndash, 10 cm soil layer but were not significantly different among forest types in the lower soil layers. There were positive correlations between these parameters and aboveground biomass across forest types in soil layer of 0&ndash, 10 cm, but not in the lower soil layers. Fine root turnover rate was generally higher in mixed forests than in monocultures at all soil depths. In conclusion, the natural regeneration procedure after clear-cutting in the subalpine region of western Sichuan seems to be superior from the perspective of fine root dynamics.

A pot experiment was conducted to investigate the effects of nitrogen (N) addition (0, 20, 40 g N m-2 year-1, N0, N20, N40, respectively) on the growth, and biomass accumulation and allocation of coniferous and deciduous (Picea asperata Mast. and Betula albosinensis Burk.) seedlings under a range of soil moisture limitation (40%, 50%, 60%, 80% and 100% of field capacity, FC). At 100% FC, growth of shade-tolerant P. asperata increased with N supply, while that of shade-intolerant B. albosinensis reached a maximum at N20, declining somewhat thereafter. At 60% FC and lower moisture content, water availability limited the growth of P. asperata seedlings, while N availability became progressively limiting to growth with moisture increasing above 60% FC. The transition from principally water-limited response to N-limited response in B. albosinensis occurred at lower moisture content. For P. asperata, these patterns reflected the responses of roots, consistent with changes in root/shoot biomass. For B. albosinensis the response reflected changes in shoot dimensions and root biomass fraction, the latter decreasing with size and foliar [N]. We are not aware of another study demonstrating such differences in the shape of the growth responses of seedlings of differing potential growth rate, across a range in belowground resource supply. The responses of leaf photosynthesis (as well as photosynthetic water and N-use efficiencies) were consistent with the observed growth response of P. asperata to water and N availability, but not of B. albosinensis, suggesting that leaf area dynamics (not measured) dominated the response of this species. Betula albosinensis, a fast-growing species, has a relative narrow range of soil water and N availability for maximum growth, achieved by preferential allocation to the shoot as resources meet the requirements at moderate N and water supply. In contrast, P. asperata increases shoot biomass progressively with increasing resources up to moderate water supply, preferentially growing more roots when resources are not limiting, suggesting that its capacity to produce shoot biomass may reach a biological limit at moderate levels of resource supply.

Soil microbial community plays a crucial role in fine roots decomposition in the forest ecosystem, and understanding the dynamics of soil microbial diversity in the fine roots will be helpful to reveal the mechanism of fine root decomposition at different critical periods. As yet, little is known about the dynamics of soil microbial diversity in the process of fine roots decomposition during the soil freeze–thaw season in the high-frigid forest ecosystem. In order to understand the effects of seasonal freeze–thaw cycles on bacterial diversity in the process of fine roots decomposition at different critical periods in cold winter, litterbags with spruce (Picea asperata), fir (Abies faxoniana) and birch (Betula albosinensis) fine roots were buried in the forest floor at altitudes of 3582 m, 3298 m, and 3023 m in different sites in the eastern Tibet Plateau on October 26, 2009. The litterbags were picked up and stored in ice box at the periods of onset of freezing (OF), deep freezing (DF), early thawing stage (ETS), middle thawing stage (MTS), late thawing stage (LTS), and early growing season (EGS) from freeze–thaw season to early growing season. DNA of bacterial in fine roots taken back was extracted immediately. Based on the PCR-DGGE technique, diverse bacterial communities in the studied fine roots were found in the whole process of fine roots decomposition. Bacterial diversity in fine roots were significantly higher at the periods of onset of freezing, middle thawing and late thawing, but significantly lower at the periods of in deep freezing and early thawing. Bacterial diversity in decomposing fine roots varied greatly with seasonal soil freeze–thaw process and tree species, but varied slightly with the studied altitudes. Regardless of tree species, higher similarity of bacterial community in fine roots was observed among different altitudes at the same period, while relatively lower similarity of bacterial community in fine roots was observed at different critical periods at the same altitude. However, the similarity of bacterial community in fine roots was always higher than 48% during the freeze–thaw season. Biodiversity indices of richness, Shannon–Wiener index, Simpson index and evenness of bacterial community were correlated significantly with average, maximum and minimum temperatures in soils during the freeze–thaw season, only except for the correlation between soil average temperature and Simpson index. The nitrogen concentration in fine roots was correlated significantly with Simpson index and evenness of bacterial community, but the phosphorus concentration was not significantly correlated with these bacterial indices. In conclusion, continuous soil freezing reduced the level of bacterial community diversity in fine roots, and the bacterial community diversity in fine roots at soil thawing period would recover the level at onset freezing period, regardless of the studied altitudes. The dynamics of bacterial community in fine roots conformed to the dynamics of root decomposition. Meanwhile, fine roots quality could also influence the level of bacterial community diversity.

Investigating the responses of trees to the heterogeneous distribution of nutrients in soil and simultaneous presence of neighboring roots could strengthen the understanding of an influential mechanism on tree growth and provide a scientific basis for forest management. Here, we conducted two split-pot experiments to investigate the effects of nutrient heterogeneity and intraspecific competition on the fine root morphology and nutrient capture of Picea asperata. The results showed that P. asperata efficiently captured nutrients by increasing the specific root length (SRL) and specific root area (SRA) of first-and second-order roots and decreasing the tissue density of first-order roots to avoid competition for resources and space with neighboring roots. The nutrient heterogeneity and addition of fertilization did not affect the fine root morphology, but enhanced the P and K concentrations in the fine roots in the absence of a competitor. On the interaction between nutrient heterogeneity and competition, competition decreased the SRL and SRA but enhanced the capture of K under heterogeneous soil compared with under homogeneous soil. Additionally, the P concentration, but not the K concentration, was linearly correlated to root morphology in heterogeneous soil, even when competition was present. The results suggested that root morphological features were only stimulated when the soil nutrients were insufficient for plant growth and the nutrients accumulations by root were mainly affected by the soil nutrients more than the root morphology.

Partial desiccation treatment (PDT) promotes the germination capacity of conifer somatic embryos. Lysine acetylation (LysAc) is a dynamic and reversible post-translational modification that plays a key role in many biological processes including metabolic pathways and stress response. To investigate the functional impact of LysAc in the response of Picea asperata somatic embryos to PDT, we performed a global lysine acetylome analysis. Here, combining antibody-based affinity enrichment and high-resolution mass spectrometry, we identified and validated 1079 acetylation sites in 556 acetylated proteins from P. asperata somatic embryos during PDT. These data represent a novel large-scale dataset of lysine-acetylated proteins from the conifer family. Intensive bioinformatics analysis of the Gene Ontology of molecular functions demonstrated that lysine-acetylated proteins were mainly associated with binding, catalytic activities, and structural molecular activities. Functional characterization of the acetylated proteins revealed that in the desiccated somatic embryos, LysAc is mainly involved in the response to stress and central metabolism. Accordingly, the majority of these interacting proteins were also highly enriched in ribosome, proteasome, spliceosome, and carbon metabolism clusters. This work provides the most comprehensive profile of LysAc for a coniferous species obtained to date and facilitates the systematic study of the physiological role of LysAc in desiccated somatic embryos of P. asperata.

Emissions of greenhouse gases (GHG) such as CO2 and N2O from soils are affected by many factors such as climate change, soil carbon content, and soil nutrient conditions. However, the response patterns and controls of soil CO2 and N2O fluxes to global warming and nitrogen (N) fertilization are still not clear in subalpine forests. To address this issue, we conducted an eight-year field experiment with warming and N fertilization treatments in a subalpine coniferous spruce (Picea asperata Mast.) plantation forest in China. Soil CO2 and N2O fluxes were measured using a static chamber method, and soils were sampled to analyze soil carbon and N contents, soil microbial substrate utilization (MSU) patterns, and microbial functional diversity. Results showed that the mean annual CO2 and N2O fluxes were 36.04 &plusmn, 3.77 mg C m&minus, 2 h&minus, 1 and 0.51 &plusmn, 0.11 &micro, g N m&minus, 1, respectively. Soil CO2 flux was only affected by warming while soil N2O flux was significantly enhanced by N fertilization and its interaction with warming. Warming enhanced dissolve organic carbon (DOC) and MSU, reduced soil organic carbon (SOC) and microbial biomass carbon (MBC), and constrained the microbial metabolic activity and microbial functional diversity, resulting in a decrease in soil CO2 emission. The analysis of structural equation model indicated that MSU had dominant direct negative effect on soil CO2 flux but had direct positive effect on soil N2O flux. DOC and MBC had indirect positive effects on soil CO2 flux while soil NH4+-N had direct negative effect on soil CO2 and N2O fluxes. This study revealed different response patterns and controlling factors of soil CO2 and N2O fluxes in the subalpine plantation forest, and highlighted the importance of soil microbial contributions to GHG fluxes under climate warming and N deposition.

The effects of soil warming and nitrogen (N) fertilization on nitrogen metabolism and nutrient status of Picea asperata Mast. and Pinus tabulaformis Carr. seedlings were studied in this paper. Infrared heaters increased monthly average soil and air temperature by 2.6 and 2.1°C above the ambient, respectively. Ammonium nitrate (NH4NO3) was added in an amount equivalent to an additional 25 g N m−2 a−1. Interestingly, soil warming, N fertilization, and their combination decreased foliar phosphorus (P) and magnesium (Mg) concentration of Picea asperata seedlings, but increased them in Pinus tabulaformis seedlings. Moreover, the combination of warming and N fertilization induced greater increments of amino acid, area-based N concentration and mass-based N concentration, manganese (Mn) and zinc (Zn), and further decreased P and calcium (Ca) concentration in Picea asperata seedlings. However, this phenomenon was not observed in Pinus tabulaformis seedlings. These results indicated that Picea asperata seedlings a...

Divergence during the early stage of speciation can be driven by a population bottleneck via reduced gene flow and enhanced lineage sorting. In this study, we aimed to examine whether such bottlenecks occurred during the initial speciation of two closely related spruce species Picea asperata and P. crassifolia occurring on the Qinghai–Tibet Plateau (QTP). We analyzed sequences of three chloroplast, two mitochondrial DNA fragments and a further 13 nuclear loci from 216 individuals of the two species. Both species showed a low level of genetic diversity in contrast to other congeners occurring in the QTP and adjacent regions. The estimated population sizes of P. asperata and P. crassifolia are less than the ancestral population size before splitting. These results together with multiple statistical tests (Tajima's D, Fu and Li's D* and F*) suggest that these two species underwent recent bottlenecks. Based on approximate Bayesian computation (ABC), we also determined that the period of the population shrinkage was consistent with the interspecific divergence during the late Pleistocene. The reduced population sizes and the divergent selection may together have triggered the initial divergence under high gene flow between these two species. Our results therefore highlight the importance of climatic oscillations during the late Pleistocene in promoting speciation through changing demographic sizes of the ancestral species on the QTP and in adjacent regions.

Tree species can exert a strong influence on rhizosphere nutrient cycling through root and rhizosphere processes and create feedback in the patterns of nutrient cycling in forest ecosystems. In this study, we conducted an experiment to compare the rhizosphere effects of two coniferous species on N transformation as well as their responses to experimental warming using infrared heaters in the Eastern Tibetan Plateau. We examined the potential net N mineralization and nitrification rates. N availability, and microbial biomass C (MBC) and N (MBN) in rhizosphere soils of Picea asperata and Abies faxoniana plots and compared them to bulk soils. The infrared heater increased both the mean air and the soil temperatures by 1.5 degrees C and 2.1 degrees C respectively. Potential net N mineralization and net nitrification rates were generally greater in rhizosphere soils for the two conifers than in bulk soil, especially in the warmed plots. This led to higher NH4+ and NO3- concentrations in the rhizosphere soils. MBC and MBN were markedly higher in the rhizosphere soils relative to bulk soil in the study plots. In the control subplots of P. asperata, MBC, MBN, potential net N mineralization and net nitrification rates in the rhizosphere were 9.6%, 21.7%, 33.3% and 20.1% greater than in the bulk soil, respectively. MBC, MBN, potential net N mineralization and net nitrification rates in the control subplots of A. faxoniana, however, were 2.0%, 7.7%, 22.0% and 11.8% higher, respectively, in the rhizosphere than in the bulk soil; all of the variables were significantly lower than those of P. asperata subplots. Warming significantly promoted N transformation and nutrient availability by enhancing the rhizosphere priming effects for the two conifers, but the magnitudes of the rhizosphere effects on soil N transformation stimulated by warming were generally greater in P. asperata than in A. faxoniana subplots. Differences in the altered morphological and functional characteristics of the roots between the two species under experimental warming could be largely responsible for this variation. Taken together, the results indicated that the two species exhibited similar patterns but with considerably different magnitudes of rhizosphere effects on N transformations in response to experimental warming, implying different capacities of the two conifers to acquire nutrients and thereby altered the competitive and adaptive relationships between the tree species under climate change. (c) 2012 Elsevier B.V. All rights reserved.

The aim of this study was to analyze the composition and spatio-temporal variation of soil macroinvertebrate communities in the northern Hengduanshan Mountains, Southwest China. The soil macroinvertebrates were investigated in April, August and November of 2008 in two deciduous broad-leaved forests: secondary shrub forest (SSF) and Betula albosinensis forest (BSF), two coniferous forests: Picea asperata plantation (PAP) and Abies fabri and Larix kaempferi mixed forest (ALF), a coniferous (A. fabri) and broad leaf (B. albosinensis) mixed forest (ABF) and a subalpine meadow (SM) at elevations of 2659 to 3845 m. The soil macroinvertebrate community showed typical zonal characteristics compared with other localities in China. It comprised at least 113 taxa and was dominated by Insects (43.33%, mainly Coleoptera and Diptera), followed by Diplopoda (16.37%), Malacostraca (14.31%), Oligochaeta (12.71%) and Chilopoda (6.67%). Abundances of each taxa showed clear differences among habitats. Community abundance ranged from 71.56 individuals/m2 in the PAP to 148.00 individuals/m2 in the SM. The composition and structure of the soil macroinvertebrate communities varied among the six habitats and changed with the sampling periods. The richness, abundance and Shannon index of soil macroinvertebrate communities were significantly different among habitats, but not between sampling periods. Significant temporal responses in abundance were recorded in the SSF and BSF and the Shannon index for the SM. Further analysis showed that the abundance of the seven taxonomic groups also differed significantly among habitats, but that sampling period only had significant effects on a few taxonomic groups. Patterns of temporal variation in abundance differed among the seven taxonomic groups, even when the same taxa also differed markedly among habitat types. On the whole, these temporal variations in abundance on community and taxa group levels were greater in the SSF and BSF than in the PAP, ABF, ALF and the SM. However, the habitat had a stronger effect on soil macroinvertebrates than the sampling period. Moreover, a significant correlation was found between the abundance, richness and Shannon index of soil macroinvertebrate community and plant species richness, but not with soil properties or elevation. Additionally, more taxonomic groups were significantly influenced by the litter mass and plant coverage, but few with soil properties. Plant community was the main environmental determinant influencing macroinvertebrate distribution in ecosystem of the northern Hengduanshan Mountains and presented a greater effect on the soil macroinvertebrates than soil properties or elevation. The temporal effects on soil macroinvertebrates were stronger in the deciduous broad-leaved forests than in the coniferous forests.

Ectomycorrhiza (ECM) plays an important role in plant nitrogen (N) nutrition and regulates plant responded to climate warming. We conducted a field experiment in a natural forest and a plantation in the eastern Tibetan Plateau to estimate the warming effects of open-top chambers (OTC) on ECM and N nutrition of Picea asperata seedlings. Four-year warming significantly decreased ECM colonization, ECM fungal biomass, fine root vigor and the N concentration of leaf, stem and coarse root, but significantly increased fine root N concentration and N content of leaf, stem, fine root and whole plant in natural forest. Contrarily, warming induced no obvious change in most of these parameters in plantation. Moreover, warming decreased rhizospheric soil inorganic N content in both forests. Our results showed that four-year warming was not beneficial for ECM colonization of P. asperata seedlings in the two forests and the seedlings in natural forest were more sensitive and flexible to experimental warming than in plantation. The changes of ECM colonization and fine root biomass for effective N uptake would be good for plant growth and remit N leaching under future warming in natural forest.

Summary Naturally regenerated seedlings or saplings in post-harvest forest stands are expected to contribute to population regeneration and the restoration and maintenance of genetic diversity in forest trees. In this study, we used intact old-growth stands of dragon spruce ( Picea asperata Mast.) as a benchmark to compare genetic diversity between mature and progeny cohorts sampled from post-harvest planted and post-harvest naturally regenerated stands in western Sichuan, China. Mature cohorts in the intact stands exhibited the highest levels of genetic diversity among the three types of stands. Genetic diversity within the progeny cohorts in the intact stands was significantly lower than their parent cohorts for all five measures of diversity. The progeny cohorts of the intact stands had lower genetic diversity than progeny cohorts from two types of post-harvest stands, but only for two measures of diversity. There were no significant differences in diversity between the mature or progeny cohorts of the planted and naturally regenerated stands. When mature and progeny cohorts were pooled, genetic variation within the united sample was enriched and close to that of the mature cohort in the intact stands. High genetic similarity and low genetic differentiation between progeny cohorts in the post-harvest stands and mature cohorts in the intact stands indicated a sufficient seed or pollen migration from the surrounding intact stands into the regenerated area. These findings illustrate the important role of the mature cohorts in the intact stands as reservoirs of genetic diversity.

The endemic Chinese Grouse (Tetrastes sewerzowi) inhabits subalpine coniferous forests. Little is known about its diet in the breeding season, especially while birds feed on the ground. Analysis of crop contents indicated that willow (Salix spp.) was the primary food of males (>98% wet weight, n = 4), whereas Dragon spruce (Picea asperata) seeds and willow were the primary foods of females (both >40% wet weight, n = 2). Dragon spruce seeds, invertebrates (mainly ants), and forbs were frequently consumed by females, but seldom by males, possibly to meet the nutrient constraints of egg formation. We suggest the different diets of males and females of this monogamous species may be the result of females allocating more time to searching for scarce, nutritious food, whereas males spend more time in vigilance behavior.

The aim of this study was to examine the response of internal conductance to CO(2) (g(i)) to soil water deficit and contrasting light conditions, and their consequences on photosynthetic physiology in two Picea asperata Mast. populations originating from wet and dry climate regions of China. Four-year-old trees were subjected to two light treatments (30% and 100% of full sunlight) and two watering regimes (well watered, drought) for 2 years. In both tested populations, drought significantly decreased g(i) and the net photosynthesis rate (A) and increased carbon isotope composition (delta(13)C) values in both light treatments, in particular in the sun. Moreover, drought resulted in a significantly higher relative limitation due to stomatal conductance (L(s)) in both light treatments and higher relative limitation due to internal conductance (L(i)) and abscisic acid (ABA) in the sun plants. The results also showed that L(i) (0.26-0.47) was always greater than L(s) (0.12-0.28). On the other hand, drought significantly decreased the ratio of chloroplastic to internal CO(2) concentration (C(c)/C(i)), photosynthetic nitrogen utilization efficiency (PNUE) and total biomass in the sun plants of the wet climate population, whereas there were no significant changes in these parameters in the dry climate population. Our results also showed that the dry climate population possessed higher delta(13)C values with higher ratio of internal conductance to stomatal conductance (g(i)/g(s)), suggesting that increasing the g(i)/g(s) ratio enhances water-use efficiency (WUE) in plants evolved in arid environments. Thus, we propose that the use of the g(i)/g(s) parameter to screen P. asperata plants with higher water deficit tolerance is certainly worthy of consideration. Furthermore, g(i) is an important variable, which reflects the population differences in PNUE, and it should thus be included in plant physiological investigations related to leaf economics.

The morphological and physiological plasticity to 6 light conditions was investigated for seedlings of 4 tree species dominating at different successional serals in subalpine coniferous forests in eastern Qinghai-Tibet Plateau, China. Abies faxoniana is a late successional species, while Betula albo-sinensis is a pioneer tree species, with Picea asperata and Acer davidii among other mid-late successional species. To compare the responses of photosynthetic characters to different growth light conditions, the seedlings were potted and placed in artificially shaded chambers with gradients of 100%, 55%, 40%, 25%, 15% and 7% of the full sunlight, respectively. During two and a half years' cultivation, various morphological and photosynthetic parameters were measured and analyzed. The results were: 1) all seedlings of the four species under the low growth light conditions showed decrease in root collar diameter, relative growth rate, leaf thickness, root mass ratio, leaf area-based photosynthetic capacity, dark respiration rate, light saturation point and light compensation point, while showed increase in specific leaf area, above-ground to under-ground mass ratio, specific stem length, leaf mass ratio and stem mass ratio; 2) under most light conditions, A. faxoniana of the two conifers revealed lower values in both leaf area-based photosynthetic capacity and respiration rates than P. asperata ; the same fact held true for the two broad-leaved trees with lower values observed in B. albo-sinensis ; 3) under higher light conditions, relative growth rates of P . asperata and B . albo-sinensis got higher values than those of A . faxoniana and A . davidii , while contrary results were obtained under lower light regimes; 4) the means of phenotypic plastic indices of the eleven morphological and physiological parameters of P . asperata and B . albo-sinensis were higher than those of A . faxoniana and A . davidii , respectively. The findings indicate that A . faxoniana has better adaptation to low light regimes, but as a shade-tolerant species, it is not so adapted to low light regimes as the trees of earlier successional serals, especially P . asperata and B . albo-sinensis . Of the four tree species, physiological plastic indices were higher than morphological plastic indices, suggesting that morphological plasticity plays an important role in their adaptation to different growth light conditions. The results also support the hypothesis that ecophysiological traits of tree species determine their successional status and associate habitats of their seedlings.

Ultraviolet-B (UV-B) radiation and nitrogen are expected to increase simultaneously with future changes in global climate. In this study, growth and photosynthetic responses of Picea asperata seedlings to enhanced UV-B and to nitrogen supply were studied. The experimental design included two levels of UV-B treatments (ambient UV-B, 11.02 kJ m-2 d-1; enhanced UV-B, 14.33 kJ m-2 d-1) and two nitrogen levels (0; 20 g N m-2) to determine whether nitrogen can alleviate the negative impacts of enhanced UV-B on seedling growth and photosynthesis. Enhanced UV-B significantly inhibited plant growth and impaired net photosynthetic rate, stomatal conductance, transpiration rate, the light-saturated assimilation rate, assimilation capacity, light compensation point, dark respiration rate, apparent quantum yield, photosynthetic pigments and maximum quantum yield of photosynthesis of P. asperata seedlings, whereas minimal fluorescence and intercellular CO2 concentration increased by enhanced UV-B. On the other hand, nitrogen supply improved the photosynthetic performance and plant growth, but only under ambient UV-B. In fact, nitrogen supply could not alleviate the photosynthetic impairments in P. asperata seedlings exposed to enhanced UV-B radiation.

The combined effects of water stress (WS) and low irradiance (LI) on growth, photosynthesis, osmotic adjustment, and lipid peroxidation were studied in dragon spruce (Picea asperata Mast.) seedlings grown under two water treatments (well watered, 100 % of field capacity, and water stressed, 30 % of field capacity) and two irradiances (HI, 100 % of full sunlight and LI, 15 % of full sunlight). WS reduced growth, chlorophyll (Chl) a and b contents, net photosynthetic rate, transpiration rate, stomatal conductance, and effective quantum yield of photosystem 2 (Y) but increased free proline and malondialdehyde contents. LI increased Chl contents and decreased Y, photochemical quenching (qP), and non-photochemical quenching (qN) under both water treatments. Hence the seedlings in the understory were more sensitive to drought than to LI.

The effects of exogenous abscisic acid (ABA) on the acclimation of Picea asperata to water deficit were investigated in two populations originating from wet and dry climate regions of China. Exogenous ABA was sprayed onto the leaves, and changes in plant growth and structure, gas exchange, water use efficiency (WUE), endogenous ABA content, and antioxidant enzyme levels were monitored. The results demonstrated that ABA application affected the two P. asperata populations in different ways during the water deficit. ABA application resulted in significantly lower CO(2) assimilation rates (A) under water deficit in plants from the wet climate population, whereas there were no significant changes in this parameter in the dry climate population. On the other hand, ABA application significantly decreased the dry shoot biomass, stomatal conductance (g(s)), transpiration rate (E), and malondialdehyde (MDA) content, and it significantly increased the leaf mass per area (LMA), root/shoot ratio (Rs), fine root/total root ratio (Ft), WUE, ABA content, and the superoxide dismutase (SOD), ascorbate peroxidase (APX), and catalase (CAT) activities under water-deficit conditions in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet climate population. The results clearly demonstrated that sensitivity to an exogenous ABA application is population-dependent in P. asperata. Direct evidence is presented that variation in physiological mechanisms rather than different rates of ABA absorption explain the population differentiation in the sensitivity to exogenous ABA, and that the physiological basis for the amplified response to water deficit caused by exogenous ABA, present mainly in the dry climate population, is related to internal ABA accumulation. These results provide evidence for adaptive differentiation between populations of P. asperata, and they support the expected relationship between environmental heterogeneity and the magnitude of plastic responses in plant populations.