Variation in photosynthetic capacity of Salvia przewalskii along elevational gradients on the eastern Qinghai-Tibetan Plateau, China.

Elevational variation in plant growing environment drives diversification of photosynthetic capacity, however, the mechanism behind this reaction is poorly understood. We measured leaf gas exchange, chlorophyll fluorescence, anatomical characteristics, and biochemical traits of Salvia przewalskii at elevations ranging from 2400 m to 3400 m above sea level (a.s.l) on the eastern Qinghai-Tibetan Plateau, China. We found that photosynthetic capacity showed an initial increase and then a decrease with rising elevation, and the best state observed at 2800 m a.s.l. Environmental factors indirectly regulated photosynthetic capacity by affecting stomatal conductance (gs), mesophyll conductance (gm), maximum velocity of carboxylation (Vc max), and maximum capacity for photosynthetic electron transport (Jmax). The average temperature (T) and total precipitation (P) during the growing season had the highest contribution to the variation of photosynthetic capacity of S. przewalskii in subalpine areas, which were 25% and 24%, respectively. Photosynthetic capacity was mainly affected by diffusional limitations (71%-89%), and mesophyll limitation (lm) played a leading role. The variation of gm was attributed to the effects of environmental factors on the volume fraction of intercellular air space (fias), the thickness of cell wall (Tcw), the surface of mesophyll cells and chloroplasts exposed to intercellular airspace (Sm, Sc), and plasma membrane intrinsic protein (PIPs, PIP1, PIP2), independent of carbonic anhydrase (CA). Optimization of leaf tissue structure and adaptive physiological responses enabled plants to efficiently cope with variable climate conditions of high-elevation areas, and the while maintaining high levels of carbon assimilation.

[Differences and drivers of leaf stable carbon and nitrogen isotope in herbs under different vegetation types on the eastern Qinghai-Tibet Plateau]. / é’è—é«˜åŽŸä¸œç¼˜ä¸åŒæ¤è¢«ç±»åž‹ä¸‹è‰æœ¬æ¤ç‰©å¶ç‰‡ç¢³æ°®ç¨³å®šåŒä½ç´ å·®å¼‚åŠå ¶é©±åŠ¨å› ç´ .

The natural abundance of stable carbon and nitrogen isotopes (δ13C and δ15N) in leaves can provide comprehensive information on the physiological and ecological processes of plants and has been widely used in ecological research. However, recent studies on leaf δ13C and δ15N have focused mainly on woody species, few studies have been conducted on herbs in different vegetation types, and their differences and driving factors are still unclear. In this study, we focused on the herbs in subalpine coniferous forests, alpine shrublands, and alpine mea-dows on the eastern Qinghai-Tibet Plateau, and investigated the differences in leaf δ13C and δ15N of herbs and the driving factors. The results showed that there were significant differences in leaf δ13C and δ15N values of herbs among different vegetation types, with the highest δ13C and δ15N values in alpine meadows, followed by alpine shrublands, and the lowest in subalpine coniferous forests. Using variation partitioning analysis, we revealed that differences in leaf δ13C and δ15N of herbs among various vegetation types were driven by both leaf functional traits and climate factors, with the contribution of leaf functional traits being relatively higher than that of climate factors. Hierarchical partitioning results indicated that mean annual temperature (MAT), chlorophyll content index, leaf nitrogen content per unit area (Narea), and leaf mass per area were the main drivers of leaf δ13C variations of herbs across different vegetation types, while the relative importance of Narea and MAT for variation in leaf δ15N of herbs was much higher than those other variables. There was a strong coupling relationship between leaf δ13C and δ15N as indicated by the result of the ordinary least squares regression. Our findings could provide new insights into understanding the key drivers of leaf δ13C and δ15N variations in herbs across different vegetation types.

Responses of leaf traits to altitude in Quercus aquifolioides and Sorbus rehderiana on the eastern edge of the Qinghai-Tibet Plateau, China.

As the most senstitive plant organs to environmental changes, leaves serve as crucial indicators of plant survival strategies. We measured the morphology, anatomical traits, gas exchange parameters, and chlorophyll fluorescence parameters of Quercus aquifolioides (evergreen broad-leaved) and Sorbus rehderiana (deciduous broad-leaved) at altitudes of 2600, 2800, 3000, 3200 and 3400 m on the eastern edge of the Qinghai-Tibet Plateau, China. We explored the similarity and difference in their responses to altitude change and the ecological adaptation strategy. The results showed that as the altitude increased, leaf dry matter content of Q. aquifolioides decreased, that of S. rehderiana increased, leaf size for both species gradually decreased, and the palisade coefficient of Q. aquifolioides showed a decreasing trend, contrasting with the increasing trend in S. rehderiana. As the altitude increased, the thickness of leaves, palisade tissue, spongy tissue, upper epidermis, and lower epidermis of both species increased significantly, with the increment of 22.4%, 4.9%, 45.1%, 23.3%, 19.6%, and 28.2%, 46.9%, 8.9%, 25.9%, 20.8% at altitude of 3400 m, respectively, compared with the altitude of 2600 m. The gas exchange and chlorophyll fluorescence parameters of S. rehderiana significantly increased with increasing altitude, while Q. aquifolioides showed the opposite trend. Leaf anatomical traits, gas exchange, and chlorophyll fluorescence parameters of both species displayed considerable plasticity. There were significant correlations among most leaf traits and between leaf traits and altitude. The survival strategy of Q. aquifolioides was more conservative in response to altitude changes, while that of S. rehderiana was more active. Both species adapted to different altitudes by adjusting their own traits.

Effects of slope aspect on soil enzyme activity and microbial nutrient limitation in subalpine region of wes-tern Sichuan, China.

Exploring the resource limitation of soil microbial metabolism is essential to understand ecosystem functions and processes. However, the spatially divergent patterns and drivers of soil microbial nutrient limitation cha-racteristics in montane ecosystems at small scales, especially at the slope aspect scale, are still unclear. In this study, we measured soil enzyme activities involved in carbon (C), nitrogen (N) and phosphorus (P) cycle and quantified the microbial nutrient limitations by enzyme stoichiometry in two representative mountain sites in subalpine region of western Sichuan, including the sunny and shady slopes with different vegetation types (shrubland and forest, respectively) in Miyaluo of Lixian County, and with the same vegetation type (shrubland) in Yakexia of Heishui County. The results showed that soil enzyme activities and their stoichiometric ratios were significantly different between slope aspects in Miyaluo, while the differences were not significant in Yakexia. The stoichiometry ratio of C-, N- and P-acquiring enzymes on the sunny slope of Miyaluo was 1:0.96:0.92, approaching the 1:1:1 ratio at the global scale, but deviated from 1:1:1 on the shady slope of Miyaluo (1:1.39:0.75) and the different slopes of Yakexia (1:1.09:1.35). There was no significant difference in vector length between slope aspects at both sites, indicating no significant effect of slope aspect on the microbial C limitation. The vector angle was significantly higher on the sunny slope (43.6°) than that on the shady slope (28.7°) in Miyaluo, suggesting that the microorganisms were mainly N-limited. Partial least squares path model showed that the vector angle was mainly directly influenced by the soil nutrient ratios. The vector angle ranged from 50.3° to 51.4°, and did not differ between slope aspects in Yakexia. Therefore, differences in vegetation types between slope aspects drove variations in soil enzyme activity and microbial nutrient limitation through soil properties. It would provide a scientific basis for predicting the spatial pattern of soil enzyme activity and microbial nutrient limitation.

Leaf traits divergence and correlations of woody plants among the three plant functional types on the eastern Qinghai-Tibetan Plateau, China.

Leaf traits are important indicators of plant life history and may vary according to plant functional type (PFT) and environmental conditions. In this study, we sampled woody plants from three PFTs (e.g., needle-leaved evergreens, NE; broad-leaved evergreens, BE; broad-leaved deciduous, BD) on the eastern Qinghai-Tibetan Plateau, and 110 species were collected across 50 sites. Here, the divergence and correlations of leaf traits in three PFTs and relationships between leaf traits and environment were studied. The results showed significant differences in leaf traits among three PFTs, with NE plants showed higher values than BE plants and BD plants for leaf thickness (LT), leaf dry matter content (LDMC), leaf dry mass per area (LMA), carbon: nitrogen ratio (C/N), and nitrogen content per unit area (Narea), except for nitrogen content per unit mass (Nmass). Although the correlations between leaf traits were similar across three PFTs, NE plants differed from BE plants and BD plants in the relationship between C/N and Narea. Compared with the mean annual precipitation (MAP), the mean annual temperature (MAT) was the main environmental factor that caused the difference in leaf traits among three PFTs. NE plants had a more conservative approach to survival compared to BE plants and BD plants. This study shed light on the regional-scale variation in leaf traits and the relationships among leaf traits, PFT, and environment. These findings have important implications for the development of regional-scale dynamic vegetation models and for understanding how plants respond and adapt to environmental change.

Contrasting responses of soil microbial biomass and extracellular enzyme activity along an elevation gradient on the eastern Qinghai-Tibetan Plateau.

Soil microbial community composition and extracellular enzyme activity are two main drivers of biogeochemical cycling. Knowledge about their elevational patterns is of great importance for predicting ecosystem functioning in response to climate change. Nevertheless, there is no consensus on how soil microbial community composition and extracellular enzyme activity vary with elevation, and little is known about their elevational variations on the eastern Qinghai-Tibetan Plateau, a region sensitive to global change. We therefore investigated the soil microbial community composition using phospholipid fatty acids (PLFAs) analysis, and enzyme activities at 2,820 m (coniferous and broadleaved mixed forest), 3,160 m (dark coniferous forest), 3,420 m (alpine dwarf forest), and 4,280 m (alpine shrubland) above sea level. Our results showed that soil microbial community composition and extracellular enzyme activities changed significantly along the elevational gradient. Biomass of total microbes, bacteria, and arbuscular mycorrhizal fungi at the highest elevation were the significantly lowest among the four elevations. In contrast, extracellular enzyme activities involved in carbon (C)-, nitrogen (N)-, and phosphorus (P)- acquiring exhibited the maximum values at the highest elevation. Total nutrients and available nutrients, especially P availability jointly explained the elevational pattern of soil microbial community, while the elevational variation of extracellular enzyme activities was dependent on total nutrients. Microbial metabolism was mainly C- and P-limited with an increasing C limitation but a decreasing P limitation along the elevational gradient, which was related significantly to mean annual temperature and total P. These results indicated a vital role of soil P in driving the elevational patterns of soil microbial community and metabolism. Overall, the study highlighted the contrasting responses of soil microbial biomass and extracellular enzyme activities to elevation, possibly suggesting the differences in adaption strategy between population growth and resource acquisition responding to elevation. The results provide essential information for understanding and predicting the response of belowground community and function to climate change on the eastern Qinghai-Tibetan Plateau.

Leaf functional traits have more contributions than climate to the variations of leaf stable carbon isotope of different plant functional types on the eastern Qinghai-Tibetan Plateau.

Elucidating the mechanisms that control the leaf stable carbon isotope values (δ13Cleaf) is the prerequisite for the widespread application of δ13Cleaf. However, the competing effects of physiological and environmental factors on δ13Cleaf variations of the different plant functional types (PFTs) have not been disentangled, and the corresponding mechanisms remain unclear. Based on large-scale δ13Cleaf measurements on the eastern Qinghai-Tibetan Plateau, the relative contributions and regulatory pathways of leaf functional traits (LFTs) and climatic factors to δ13Cleaf variations of the different PFTs were investigated. We found that δ13Cleaf of the different PFTs was correlated with annual mean precipitation negatively, but not a simple linear relationship with annual mean temperature and varied by PFTs. Leaf nitrogen content per unit area and leaf mass per area (correlated with δ13Cleaf positively) had more substantial effects on the δ13Cleaf variations of the different PFTs than other LFTs. The relative contributions of LFTs to the δ13Cleaf variations were greater than that of climatic factors, and the direct and indirect effects of climatic factors on δ13Cleaf variations varied by PFTs. Our findings provide new insights into understanding key drivers of δ13Cleaf variations at the PFT level on a regional scale.

The below-ground carbon and nitrogen cycling patterns of different mycorrhizal forests on the eastern Qinghai-Tibetan Plateau.

Mycorrhizal fungi can form symbiotic associations with tree species, which not only play an important role in plant survival and growth, but also in soil carbon (C) and nitrogen (N) cycling. However, the understanding of differences in soil C and N cycling patterns among forests with different mycorrhizal types is still incomplete. In order to determine the similarities and differences of soil C and N cycling patterns in different mycorrhizal forest types, three primary forests dominated by ectomycorrhizal (EcM), arbuscular mycorrhizal (AM) and ericoid mycorrhizal (ErM) trees respectively were studied on the eastern Qinghai-Tibetan Plateau. Indicators associated with soil C and N cycling, including leaf litter quality, soil C and N contents, soil C and N fluxes, and soil microbial biomass C and N contents were measured in each mycorrhizal type forest. The results showed that leaf litter quality was significantly lower with high C:N ratio and lignin: N ratio in ErM forest than that in AM and EcM forests. Soil CO2 flux (508.25 ± 65.51 mg m-2 h-1) in AM forest was significantly higher than that in EcM forest (387.18 ± 56.19 mg m-2 h-1) and ErM forest (177.87 ± 58.40 mg m-2 h-1). Furthermore, soil inorganic N content was higher in the AM forest than that in EcM and ErM forests. Soil net N mineralization rate (-0.02 ± 0.03 mg kg-1 d-1) was lower in ErM forest than that in EcM and AM forests. We speculated that AM and EcM forests were relatively characterized by rapid soil C cycling comparing to ErM forest. The soil N cycling in EcM and ErM forests were lower, implying they were 'organic' N nutrition patterns, and the pattern in ErM forest was more obvious.

Altitudinal Variation Influences Soil Fungal Community Composition and Diversity in Alpine-Gorge Region on the Eastern Qinghai-Tibetan Plateau.

Soil fungi play an integral and essential role in maintaining soil ecosystem functions. The understanding of altitude variations and their drivers of soil fungal community composition and diversity remains relatively unclear. Mountains provide an open, natural platform for studying how the soil fungal community responds to climatic variability at a short altitude distance. Using the Illumina MiSeq high-throughput sequencing technique, we examined soil fungal community composition and diversity among seven vegetation types (dry valley shrub, valley-mountain ecotone broadleaved mixed forest, subalpine broadleaved mixed forest, subalpine coniferous-broadleaved mixed forest, subalpine coniferous forest, alpine shrub meadow, alpine meadow) along a 2582 m altitude gradient in the alpine-gorge region on the eastern Qinghai-Tibetan Plateau. Ascomycota (47.72%), Basidiomycota (36.58%), and Mortierellomycota (12.14%) were the top three soil fungal dominant phyla in all samples. Soil fungal community composition differed significantly among the seven vegetation types along altitude gradients. The α-diversity of soil total fungi and symbiotic fungi had a distinct hollow pattern, while saprophytic fungi and pathogenic fungi showed no obvious pattern along altitude gradients. The ß-diversity of soil total fungi, symbiotic fungi, saprophytic fungi, and pathogenic fungi was derived mainly from species turnover processes and exhibited a significant altitude distance-decay pattern. Soil properties explained 31.27-34.91% of variation in soil fungal (total and trophic modes) community composition along altitude gradients, and the effects of soil nutrients on fungal community composition varied by trophic modes. Soil pH was the main factor affecting α-diversity of soil fungi along altitude gradients. The ß-diversity and turnover components of soil total fungi and saprophytic fungi were affected by soil properties and geographic distance, while those of symbiotic fungi and pathogenic fungi were affected only by soil properties. This study deepens our knowledge regarding altitude variations and their drivers of soil fungal community composition and diversity, and confirms that the effects of soil properties on soil fungal community composition and diversity vary by trophic modes along altitude gradients in the alpine-gorge region.

Does Forest Soil Fungal Community Respond to Short-Term Simulated Nitrogen Deposition in Different Forests in Eastern China?

Nitrogen (N) deposition has changed plants and soil microbes remarkably, which deeply alters the structures and functions of terrestrial ecosystems. However, how forest fungal diversity, community compositions, and their potential functions respond to N deposition is still lacking in exploration at a large scale. In this study, we conducted a short-term (4-5 years) experiment of artificial N addition to simulated N deposition in five typical forest ecosystems across eastern China, which includes tropical montane rainforest, subtropical evergreen broadleaved forest, temperate deciduous broadleaved forest, temperate broadleaved and conifer mixed forest, and boreal forest along a latitudinal gradient from tropical to cold temperature zones. Fungal compositions were identified using high-throughput sequencing at the topsoil layer. The results showed that fungal diversity and fungal community compositions among forests varied apparently for both unfertilized and fertilized soils. Generally, soil fungal diversity, communities, and their potential functions responded sluggishly to short-term N addition, whereas the fungal Shannon index was increased in the tropical forest. In addition, environmental heterogeneity explained most of the variation among fungal communities along the latitudinal gradient. Specifically, soil C: N ratio and soil water content were the most important factors driving fungal diversity, whereas mean annual temperature and microbial nutrient limitation mainly shaped fungal community structure and functional compositions. Topsoil fungal communities in eastern forest ecosystems in China were more sensitive to environmental heterogeneity rather than short-term N addition. Our study further emphasized the importance of simultaneously evaluating soil fungal communities in different forest types in response to atmospheric N deposition.

The effects of ectomycorrhizal and saprotropic fungi on soil nitrogen mineralization differ from those of arbuscular and ericoid mycorrhizal fungi on the eastern Qinghai-Tibetan Plateau.

Interactions between soil fungi and soil environmental factors regulate soil nitrogen (N) mineralization rates on the eastern Qinghai-Tibetan Plateau. Some studies have also illuminated differences in soil N mineralization rate based on different mycorrhizal forests, but the associated effect of soil fungal functional guilds and soil environmental factors underlying this process are not well-understood. Three primary forests respectively dominated by Abies fargesii var. faxoniana (ectomycorrhizal, EcM), Cupressus chengiana (arbuscular mycorrhizal, AM) and Rhododendron phaeochrysum (ericoid mycorrhizal, ErM) trees were selected in this area. Meanwhile, soil net N mineralization rate, soil fungal composition and soil enzyme activity among these three mycorrhizal forests were studied. Our results showed that there were significant differences in the seasonal variation of soil net N mineralization rates among three mycorrhizal forests. Soil net N mineralization rate in the AM forest was faster. EcM fungi and saprotroph are the main functional guilds in these three mycorrhizal forests. Meanwhile, the relative abundances of soil fungal functional guilds, soil temperature and soil peroxidase activity could explain 85.0% in the difference of soil net ammonification rate among three mycorrhizal forests. In addition, soil temperature, soil water-filled pore space and soil ammonium content play a central role in controlling the differing soil net nitrification rate among three mycorrhizal forests. Our results suggest differences in soil net mineralization among different mycorrhizal forest types are driven mainly by soil net ammonification. Soil fungal functional guilds and temperature regulate the rate of soil net ammonification by modulating soil peroxidase activity.

[Vertical distribution of natural abundance of stable carbon and nitrogen isotopes along the soil profile and the underlying mechanisms]. / åœŸå£¤å‰–é¢ç¢³æ°®ç¨³å®šåŒä½ç´ è‡ªç„¶ä¸°åº¦çš„åž‚ç›´åˆ†å¸ƒæ¨¡å¼åŠå ¶å½±å“æœºåˆ¶.

Understanding the changes of natural abundance of stable carbon and nitrogen isotopes (δ13C and δ15N) along soil profile is of great importance in revealing the mechanisms of soil carbon and nitrogen cycling in terrestrial ecosystems. Based on a comprehensive review on the distribution of δ13C and δ15N along soil profile, the mechanisms underlying their vertical distribution were mainly introduced here. There were three mechanisms driving the δ13C vertical distribution in soil profile: 1) historical changes of vegetation δ13C value, 2) changes of C3-C4 species dominance in plant communities, 3) accumulation of 13C-enriched microbial-derived carbon during decomposition. The effects of 13C Suess effect on the vertical distribution of δ13C in soil profile were also discussed. There were four mechanisms underlying the vertical distribution of δ15N in soil profile: 1) 15N-depletion gas loss during denitrification, 2) accumulation of 15N-enriched microbial-derived nitrogen during decomposition, 3) accumulation of 15N-encriched mycorrhizal fungi residues in deep soil as a result of transferring 15N-depleted nitrogen compounds to plants by mycorrhizae, 4) intera-ction between soil organic matter and mineral substance. We proposed important concerning points for the future study on vertical distribution of natural abundance of stable carbon and nitrogen isotopes in soil profile.

Impacts of growth form and phylogenetic relatedness on seed germination: A large-scale analysis of a subtropical regional flora.

Plant regeneration strategy plays a critical role in species survival and can be used as a proxy for the evolutionary response of species to climate change. However, information on the effects of key plant traits and phylogenetic relatedness on seed germination is limited at large regional scales that vary in climate. To test the hypotheses that phylogenetic niche conservatism plays a critical force in shaping seed ecophysiological traits across species, and also drives their response to climatic fluctuation, we conducted a controlled experiment on seed germination and determined the percentage and rate of germination for 249 species in subtropical China under two temperature regimes (i.e., daily 25°C; daily alternating 25/15°C for each 12 hr). Germination was low with a skewed distribution (mean = 38.9% at 25°C, and 43.3% at 25/15°C). One fifth of the species had low (<10%) and slow (4-30 days) germination, and only a few (8%) species had a high (>80%) and rapid (1.2-6.6 days) germination. All studied plant traits (including germination responses) showed a significant phylogenetic signal, with an exception of seed germination percentage under the alternating temperature scenario. Generalized linear models (GLMs) and phylogenetic generalized estimation equations (GEEs) demonstrated that growth form and seed dispersal mode were strong drivers of germination. Our experimental study highlights that integrating plant key traits and phylogeny is critical to predicting seed germination response to future climate change.

Fagaceae tree species allocate higher fraction of nitrogen to photosynthetic apparatus than Leguminosae in Jianfengling tropical montane rain forest, China.

Variation in photosynthetic-nitrogen use efficiency (PNUE) is generally affected by several factors such as leaf nitrogen allocation and leaf diffusional conductances to CO2, although it is still unclear which factors significantly affect PNUE in tropical montane rain forest trees. In this study, comparison of PNUE, photosynthetic capacity, leaf nitrogen allocation, and diffusional conductances to CO2 between five Fagaceae tree species and five Leguminosae tree species were analyzed in Jianfengling tropical montane rain forest, Hainan Island, China. The result showed that PNUE of Fagaceae was significantly higher than that of Leguminosae (+35.5%), attributed to lower leaf nitrogen content per area (Narea, -29.4%). The difference in nitrogen allocation was the main biochemical factor that influenced interspecific variation in PNUE of these tree species. Fagaceae species allocated a higher fraction of leaf nitrogen to the photosynthetic apparatus (PP, +43.8%), especially to Rubisco (PR, +50.0%) and bioenergetics (PB +33.3%) in comparison with Leguminosae species. Leaf mass per area (LMA) of Leguminosae species was lower than that of Fagaceae species (-15.4%). While there was no significant difference shown for mesophyll conductance (gm), Fagaceae tree species may have greater chloroplast to total leaf surface area ratios and that offset the action of thicker cell walls on gm. Furthermore, weak negative relationship between nitrogen allocation in cell walls and in Rubisco was found for Castanopsis hystrix, Cyclobalanopsis phanera and Cy. patelliformis, which might imply that nitrogen in the leaves was insufficient for both Rubisco and cell walls. In summary, our study concluded that higher PNUE might contribute to the dominance of most Fagaceae tree species in Jianfengling tropical montane rain forest.

Grazing exclusion reduced soil respiration but increased its temperature sensitivity in a Meadow Grassland on the Tibetan Plateau.

Understanding anthropogenic influences on soil respiration (R s) is critical for accurate predictions of soil carbon fluxes, but it is not known how R s responds to grazing exclusion (GE). Here, we conducted a manipulative experiment in a meadow grassland on the Tibetan Plateau to investigate the effects of GE on R s. The exclusion of livestock significantly increased soil moisture and above-ground biomass, but it decreased soil temperature, microbial biomass carbon (MBC), and R s. Regression analysis indicated that the effects of GE on R s were mainly due to changes in soil temperature, soil moisture, and MBC. Compared with the grazed blocks, GE significantly decreased soil carbon release by 23.6% over the growing season and 21.4% annually, but it increased the temperature sensitivity (Q10) of R s by 6.5% and 14.2% for the growing season and annually respectively. Therefore, GE may reduce the release of soil carbon from the Tibetan Plateau, but under future climate warming scenarios, the increases in Q10 induced by GE could lead to increased carbon emissions.