Alterations in leaf nitrogen metabolism indicated the structural changes of subtropical forest by canopy addition of nitrogen.

Globally, nitrogen deposition increment has caused forest structural changes due to imbalanced plant nitrogen metabolism and subsequent carbon assimilation. Here, a 2 consecutive-year experiment was conducted to reveal the effects of canopy addition of nitrogen (CAN) on nitrogen absorption, assimilation, and allocation in leaves of three subtropical forest woody species (Castanea henryi, Ardisia quinquegona, and Blastus cochinchinensis). We hypothesized that CAN altered leaf nitrogen absorption, assimilation and partitioning of different plants in different ways in subtropical forest. It shows that CAN increased maximum photosynthetic rate (Amax), photosynthetic nitrogen use efficiency (PNUE), and metabolic protein content of the two understory species A. quinquegona and B. cochinchinensis. By contrary, for the overstory species, C. henryi, Amax, PNUE, and metabolic protein content were significantly reduced in response to CAN. We found that changes in leaf nitrogen metabolism were mainly due to the differences in enzyme (e.g. Ribulose-1,5-bisphosphate carboxylase, nitrate reductase, nitrite reductase and glutamine synthetase) activities under CAN treatment. Our results indicated that C. henryi may be more susceptible to CAN treatment, and both A. quinquegona and B. cochinchinensis could better adapt to CAN treatment but in different ways. Our findings may partially explain the ongoing degradation of subtropical forest into a community dominated by small trees and shrubs in recent decades. It is possible that persistent high levels of atmospheric nitrogen deposition will lead to the steady replacement of dominant woody species in this subtropical forest.

Leaf hydraulic acclimation to nitrogen addition of two dominant tree species in a subtropical forest.

Plant hydraulic traits have been shown to be sensitive to changes in nitrogen (N) availability in short-term studies largely using seedlings or saplings. The extent and the magnitude of N-sensitivity of the field grown mature trees in long-term experiments, however, are relatively unknown. Here, we investigated responses of leaf water relations and morphological and anatomical traits of two dominant tree species (Castanopsis chinensis and Schima superba) to a six-year canopy N addition in a subtropical forest. We found that N addition increased leaf hydraulic conductivity in both species along with higher transpiration rate and less negative water potential at 50% loss of leaf hydraulic conductivity and at leaf turgor loss point. Examination of leaf morphological and anatomical traits revealed that increased leaf hydraulic efficiency was at least in part due to increased vessel diameter which also compromised the hydraulic safety under increased water stress. Moreover, reduced vessel reinforcement and increased thickness shrinkage index further interpreted the increases in leaf hydraulic vulnerability under N addition. Our results demonstrated that N deposition may lead to increases of plant water loss to the atmosphere as well as tree vulnerability to drought.

Dietary flexibility aids Asian earthworm invasion in North American forests.

On a local scale, invasiveness of introduced species and invasibility of habitats together determine invasion success. A key issue in invasion ecology has been how to quantify the contribution of species invasiveness and habitat invasibility separately. Conventional approaches, such as comparing the differences in traits and/or impacts of species between native and/or invaded ranges, do not determine the extent to which the performance of invaders is due to either the effects of species traits or habitat characteristics. Here we explore the interaction between two of the most widespread earthworm invaders in the world (Asian Amynthas agrestis and European Lumbricus rubellus) and study the effects of species invasiveness and habitat invasibility separately through an alternative approach of "third habitat" in Tennessee, USA. We propose that feeding behaviors of earthworms will be critical to invasion success because trophic ecology of invasive animals plays a key role in the invasion process. We found that (1) the biomass and isotopic abundances (delta13C and delta15N) of A. agrestis were not impacted by either direct effects of L. rubellus competition or indirect effects of L. rubellus-preconditioned habitat; (2) A. agrestis disrupted the relationship between L. rubellus and soil microorganisms and consequently hindered litter consumption by L. rubellus; and (3) compared to L. rubellus, A. agrestis shifted its diet more readily to consume more litter, more soil gram-positive (G+) bacteria (which may be important for litter digestion), and more non-microbial soil fauna when soil microorganisms were depleted. In conclusion, A. agrestis showed strong invasiveness through its dietary flexibility through diet shifting and superior feeding behavior and its indirectly negative effect of habitat invasibility on L. rubellus via changes in the soil microorganism community. In such context, our results expand on the resource fluctuation hypothesis and support the superior competitor hypothesis. This work presents additional approaches in invasion ecology, provides some new dimensions for further research, and contributes to a greater understanding of the importance of interactions between multiple invading species.

Disentangling the Contributions of Plant Taxonomic and Functional Diversities in Shaping Aboveground Biomass of a Restored Forest Landscape in Southern China.

Restoration is essential for supporting key ecosystem functions such as aboveground biomass production. However, the relative importance of functional versus taxonomic diversity in predicting aboveground biomass during restoration is poorly studied. Here, we used a trait-based approach to test for the importance of multiple plant diversity attributes in regulating aboveground biomass in a 30-years-old restored subtropical forest in southern China. We show that both taxonomic and functional diversities are significant and positive regulators of aboveground biomass; however, functional diversity (FD) was more important than taxonomic diversity (species richness) in controlling aboveground biomass. FD had the strongest direct effect on aboveground biomass compared with species richness, soil nutrients, and community weighted mean (CWM) traits. Our results further indicate that leaf and root morphological traits and traits related to the nutrient content in plant tissues represent the existence of a leaf and root economic spectrum, and the acquisitive resource use strategy influenced aboveground biomass. Our results suggest that both taxonomic and FD play a role in shaping aboveground biomass, but FD is more important in supporting aboveground biomass in this type of environments. These results imply that enhancing FD is important to restoring and managing degraded forest landscapes.

Plant Taxonomic Diversity Better Explains Soil Fungal and Bacterial Diversity than Functional Diversity in Restored Forest Ecosystems.

Plant attributes have direct and indirect effects on soil microbes via plant inputs and plant-mediated soil changes. However, whether plant taxonomic and functional diversities can explain the soil microbial diversity of restored forest ecosystems remains elusive. Here, we tested the linkage between plant attributes and soil microbial communities in four restored forests (Acacia species, Eucalyptus species, mixed coniferous species, mixed native species). The trait-based approaches were applied for plant properties and high-throughput Illumina sequencing was applied for fungal and bacterial diversity. The total number of soil microbial operational taxonomic units (OTUs) varied among the four forests. The highest richness of fungal OTUs was found in the Acacia forest. However, bacterial OTUs were highest in the Eucalyptus forest. Species richness was positively and significantly related to fungal and bacterial richness. Plant taxonomic diversity (species richness and species diversity) explained more of the soil microbial diversity than the functional diversity and soil properties. Prediction of fungal richness was better than that of bacterial richness. In addition, root traits explained more variation than the leaf traits. Overall, plant taxonomic diversity played a more important role than plant functional diversity and soil properties in shaping the soil microbial diversity of the four forests.

An alternative approach to reduce algorithm-derived biases in monitoring soil organic carbon changes.

Quantifying soil organic carbon (SOC) changes is a fundamental issue in ecology and sustainable agriculture. However, the algorithm-derived biases in comparing SOC status have not been fully addressed. Although the methods based on equivalent soil mass (ESM) and mineral-matter mass (EMMM) reduced biases of the conventional methods based on equivalent soil volume (ESV), they face challenges in ensuring both data comparability and accuracy of SOC estimation due to unequal basis for comparison and using unconserved reference systems. We introduce the basal mineral-matter reference systems (soils at time zero with natural porosity but no organic matter) and develop an approach based on equivalent mineral-matter volume (EMMV). To show the temporal bias, SOC change rates were recalculated with the ESV method and modified methods that referenced to soils at time t1 (ESM, EMMM, and EMMV-t1) or referenced to soils at time zero (EMMV-t0) using two datasets with contrasting SOC status. To show the spatial bias, the ESV- and EMMV-t0-derived SOC stocks were compared using datasets from six sites across biomes. We found that, in the relatively C-rich forests, SOC accumulation rates derived from the modified methods that referenced to t1 soils and from the EMMV-t0 method were 5.7%-13.6% and 20.6% higher than that calculated by the ESV method, respectively. Nevertheless, in the C-poor lands, no significant algorithmic biases of SOC estimation were observed. Finally, both the SOC stock discrepancies (ESV vs. EMMV-t0) and the proportions of this unaccounted SOC were large and site-dependent. These results suggest that although the modified methods that referenced to t1 soils could reduce the biases derived from soil volume changes, they may not properly quantify SOC changes due to using unconserved reference systems. The EMMV-t0 method provides an approach to address the two problems and is potentially useful since it enables SOC comparability and integrating SOC datasets.

Invariant community structure of soil bacteria in subtropical coniferous and broadleaved forests.

Soil bacteria may be influenced by vegetation and play important roles in global carbon efflux and nutrient cycling under global changes. Coniferous and broadleaved forests are two phyletically distinct vegetation types. Soil microbial communities in these forests have been extensively investigated but few studies have presented comparable data regarding the characteristics of bacterial communities in subtropical forests. We investigated soil bacterial biomass and community composition in three pairs of coniferous and broadleaved forests across a subtropical climatic gradient. We found that bacterial biomass differed between the coniferous and broadleaved forests across the subtropical climate gradient; however, this difference disappeared at some individual sites. In contrast, the same 90 bacterial genera were found in both forest types, and their relative abundances didn't differ between the forest types, with the exception of one genus that was more abundant in broadleaved forests. Soil nitrogen or moisture was associated with bacterial groups in the coniferous and broadleaved forests, respectively. Thus, we inferred that these forests can respond differently to future changes in nitrogen deposition or precipitation. This study highlights soil bacterial invariant community composition in contrasting subtropical forests and provides a new perspective on the potential response and feedback of forests to global changes.

Consistent effects of canopy vs. understory nitrogen addition on the soil exchangeable cations and microbial community in two contrasting forests.

Anthropogenic N deposition has been well documented to cause substantial impacts on the chemical and biological properties of forest soils. In most studies, however, atmospheric N deposition has been simulated by directly adding N to the forest floor. Such studies thus ignored the potentially significant effect of some key processes occurring in forest canopy (i.e., nitrogen retention) and may therefore have incorrectly assessed the effects of N deposition on soils. Here, we conducted an experiment that included both understory addition of N (UAN) and canopy addition of N (CAN) in two contrasting forests (temperate deciduous forest vs. subtropical evergreen forest). The goal was to determine whether the effects on soil exchangeable cations and microbial biomass differed between CAN and UAN. We found that N addition reduced pH, BS (base saturation) and exchangeable Ca and increased exchangeable Al significantly only at the temperate JGS site, and reduced the biomass of most soil microbial groups only at the subtropical SMT site. Except for soil exchangeable Mn, however, effects on soil chemical properties and soil microbial community did not significantly differ between CAN and UAN. Although biotic and abiotic soil characteristics differ significantly and the responses of both soil exchangeable cations and microbial biomass were different between the two study sites, we found no significant interactive effects between study site and N treatment approach on almost all soil properties involved in this study. In addition, N addition rate (25 vs. 50 kg N ha(-1) yr(-1)) did not show different effects on soil properties under both N addition approaches. These findings did not support previous prediction which expected that, by bypassing canopy effects (i.e., canopy retention and foliage fertilization), understory addition of N would overestimate the effects of N deposition on forest soil properties, at least for short time scale.

Ecophysiological and foliar nitrogen concentration responses of understorey Acacia spp. and Eucalyptus sp. to prescribed burning.

Eucalyptus spp. is a dominant tree genus in Australia and most Eucalyptus spp. are canopy dominant species. In Australian natural forests, Eucalyptus spp. commonly are associated with understorey legumes which play a crucial role for ecological restoration owing to their nitrogen (N) fixing ability for replenishing the soil N lost after frequent prescribed burning. This study aimed to explore to what extent physiological responses of these species differ 7 and 12 years after last fire. Two most common understorey Acacia spp., Acacia leiocalyx and A. disparrima, as well as one non-leguminous Eucalyptus resinifera, were studied due to their dominance in the forest. Both A. leiocalyx and A. disparrima showed higher carbon (C) assimilation capacity, maximum photosynthetic capacity, and moderate foliar C/N ratio compared with E. resinifera. A. leiocalyx showed various advantages compared to A. disparrima such as higher photosynthetic capacity, adaptation to wider light range and higher foliar total N (TNmass). A. leiocalyx also relied on N2-fixing ability for longer time compared to A. disparrima. The results suggested that the two Acacia spp. were more beneficial to C and N cycles for the post burning ecosystem than the non-N2-fixing species E. resinifera. A. leiocalyx had greater contribution to complementing soil N cycle long after burning compared to A. disparrima.

CAN Canopy Addition of Nitrogen Better Illustrate the Effect of Atmospheric Nitrogen Deposition on Forest Ecosystem?

Increasing atmospheric nitrogen (N) deposition could profoundly impact community structure and ecosystem functions in forests. However, conventional experiments with understory addition of N (UAN) largely neglect canopy-associated biota and processes and therefore may not realistically simulate atmospheric N deposition to generate reliable impacts on forest ecosystems. Here we, for the first time, designed a novel experiment with canopy addition of N (CAN) vs. UAN and reviewed the merits and pitfalls of the two approaches. The following hypotheses will be tested: i) UAN overestimates the N addition effects on understory and soil processes but underestimates those on canopy-associated biota and processes, ii) with low-level N addition, CAN favors canopy tree species and canopy-dwelling biota and promotes the detritus food web, and iii) with high-level N addition, CAN suppresses canopy tree species and other biota and favors rhizosphere food web. As a long-term comprehensive program, this experiment will provide opportunities for multidisciplinary collaborations, including biogeochemistry, microbiology, zoology, and plant science to examine forest ecosystem responses to atmospheric N deposition.

[Caloric value and ash content of dominant plants in plantation communities in Heshan of Guangdong, China].

Different parts of twenty dominant plant species in five plantation communities on the subtropical hilly lands in Heshan of Gunagdong as well as the litters from three of the five plantation communities were sampled, and their gross caloric value (GCV) and ash content were measured by using a PARR-1281 oxygen bomb calorimeter and a muffle furnace. Based on the measurements, the ash-free caloric value (AFCV) of the samples was calculated, and the characteristics of caloric value and ash content of the samples, according to plant part, individual, and plant growth form, were analyzed. The results showed that the GCV and AFCV of leaf, branch, stem wood, stem bark, and root were in the range of 10.7-22.17 kJ x g(-1) and 13.89-23.04 kJ x g(-1), respectively. The GCV and AFCV of leaf were significantly higher than those of other parts (P < 0.05), and the individual plant' s weighted mean values of GCV and AFCV were in the range of 14.24-19.43 and 16.63-20.99 kJ x g(-1), respectively. The mean AFCV of plantation communities was in the order of tree layer (19.55 kJ x g(-1)) > shrub layer (19.46 kJ x g(-1) > herb layer (18.77 kJ x g(-1)), with indigenous coniferous tree (19.86 kJ x g(-1)) > indigenous broad-leaved tree (19.55 kJ x g(-1)) > exotic eucalyptus (19.18 kJ x g(-1)), while the mean ash content was just the opposite. In Acacia mangium, coniferous, and Schima plantation communities, the GCV and AFCV of litters were higher than those of various plant parts (P < 0.01). The litter-falls in A. mangium and coniferous plantations had higher mean GCV and AFCV than the litters and fresh leaves of tree layer, while the fresh leaves of tree layer in Schima plantation showed higher mean GCV and AFCV.

[Time lag effect between stem sap flow and photosynthetically active radiation, vapor pressure deficit of Acacia mangium].

Based on the measurement of the stem sap flow of Acacia mangium with Granier' s thermal dissipation probe, and the cross-correlation and time serial analysis of the sap flow and corresponding photosynthetically active radiation and vapor pressure deficit, this paper studied the time lag effect between the stem sap flow of A. mangium and the driving factors of the tree canopy transpiration. The results indicated that the main driving factors of the transpiration were photosynthetically active radiation (PAR) and vapor pressure deficit (VPD). Sap flux density (Js) was more dependent on PAR than on VPD, and the dependence was more significant in dry season than in wet season. Sap flow lagged behind PAR but advanced than VPD in both dry and wet seasons. The time lag did not show any significant variation across different size tree individuals, but showed significant variation in different seasons. Time lag effect was not correlated with tree height, diameter at the breast, and canopy size. The time lag between Js and VPD was significantly related to nighttime water recharge in dry season, but reversed in wet season.

[Characteristics of CO2, CH4 and N2O emissions from winter-fallowed paddy fields in hilly area of South China].

With closed static chamber and modified gas chromatograph (HP5890 II), the in situ measurements were made on the CO2, CH4 and N2O emissions from winter-fallowed paddy fields in the hilly area of South China. Gas samples were taken simultaneously from the fields with and without rice stubble. The results showed that both of the fields had the peak value of CO2 flux in the late afternoon. In the fields with and without rice stubble, the CH4 flux was positive in the day time while negative in the night, and the N2O flux in the day time was 1.79 and 1.58 times as much as that in the night, respectively. The diurnal average CO2 flux in the field with rice stubble was significantly higher than that in bare field (P < 0.05). Correlation analysis demonstrated that the CO2 flux in winter-fallowed paddy fields had significant correlations with soil temperature, aboveground temperature, and air temperature, suggesting that temperature was the main factor affecting the CO2 emission from rice field after harvesting. During the observation time (from 2003-11-10 to 2004-01-18), the average CO2, CH4 and N2O fluxes in the field with rice stubble were (180.69 +/- 21.21) mg x m(-2) x h(-1), (-0.04 +/- 0.01) mg x m(-2) x h(-1) and (21.26 +/- 19.31) microg x m(-2) x h(-1), respectively. Compared with bare field, the CO2 flux in the field with rice stubble was 13.06% higher, CH4 absorption increased by 50%, while N2O flux was 60.75% lower. It was concluded that the winter fallowed paddy field in hilly area of South China was the source of atmospheric CO2 and N2O, and the sink of atmospheric CH4.

[Responses of canopy stomatal conductance of Acacia mangium forest to environmental driving factors].

Employing Graniers probes, this paper measured the sap flow of 14 sample trees in an Acacia mangium forest on the Heshan hilly lands of Guangdong Province, and recorded the photosynthetic active radiation (PAR), air relative humidity (RH) , and air temperature (T) above the forest canopy. The whole-tree transpiration (E), stand transpiration (Et), and mean canopy stomatal conductance (gc) were calculated, and the relationships between tree morphological characters and whole-tree water use as well as the responses of gc to PAR and vapor pressure deficit (D) were analyzed. The results showed that the whole-tree transpiration had logarithmical positive correlations with tree diameter at breast height (DBH) (P < 0.0001) , sapwood area (P < 0.0001) and canopy size (P = 0.0007), and an exponential positive correlation with tree height (P = 0. 014). The maximum gc (gc max) changed with PAR hyperbolically (P < 0.0001), and with D logarithmically (P < 0.0001). The sap flow measurement system used in this study was reliable and accurate in estimating the transpiration of whole-tree and stand and the canopy stomatal conductance, being an effective tool in studying the relationships between forest water use and environmental factors.

[Effects of nitrate application on alleviating photosynthesis restriction of Cinnamomum burmannii leaves under elevated CO2 concentration and enhanced temperature].

In this study, potted C. burmannii saplings were cultured in a top-closed chamber with elevated CO2 (+ CO2, 731 micromol x mol(-1)) and ambient CO2(CO2, 365 micromol x mol(-1)), and at diurnal temperature (day/night) 25/23 degrees C and 32/25 degrees C, respectively. The gas exchange, calculated photosynthesis parameter, Rubisco content, and activated state of Rubisco were examined. The results showed that under + CO2 and at 25/23 degrees C, the mean photosynthetic rate (Pnsat) of sapling leaves was 5.1% higher than that under + CO2 and at 32/25 degrees C. Temperature enhancement declined Pnsat, while nitrate addition increased it. Under + CO2, saplings had lower V(cmax) and J(max) at 32/25 degrees C than at 25/23 degrees C. Temperature enhancement under + CO2 also declined V(cmax) and J(max). Under + CO2, lower photorespiration rate (Rp) occurred in leaves, but temperature enhancement could increase Rp. Under + CO2, Rubisco content (NR) and its active site per unit leaf area (M) decreased as diurnal temperature changed from 25/23 degrees C to 32/25 degrees C. Meanwhile, nitrate addition could increase NR and M. It may be suggested that nitrate addition could alleviate the restriction of photosynthesis under elevated CO2 concentration and enhanced temperature.