Soil organic carbon contents,aggregate stability,and humic acid composition in different alpine grasslands in Qinghai-Tibet Plateau

Alpine grassland soils on Qinghai-Tibet Plateau store approximately 33.5 Pg of organic carbon(C) at 0–0.75 m depth and play an important role in the global carbon cycle.We investigated soil organic C(SOC),water-soluble organic C(WSOC),easily oxidizable organic C(EOC),humic C fractions,aggregate-associated C,aggregate stability,and humic acid(HA) composition along an east-west transect across Qinghai-Tibet Plateau,and explored their spatial patterns and controlling factors.The contents of SOC,WSOC,EOC,humic C fractions and aggregate-associated C,the proportions of macroaggregates(2-0.25) and micro-aggregates(0.25-0.053 mm),and the aggregate stability indices all increased in the order alpine desert alpine steppe alpine meadow.The alkyl C,O-alkyl C,and aliphatic C/aromatic C ratio of HA increased as alpine desert alpine meadow alpine steppe,and the trends were reverse for the aromatic C and HB/HI ratio.Mean annual precipitation and aboveground biomass weresignificantly correlated with the contents of SOC and its fractions,the proportions of macro- and microaggregates,and the aggregate stability indices along this transect.Among all these C fractions,SOC content and aggregate stability were more closely associated with humic C and silt and clay sized C in comparison with WSOC,EOC,and macro- and microaggregate C.The results suggested that alpine meadow soils containing higher SOC exhibited high soil aggregation and aggregate stability.Mean annual precipitation should be the main climate factor controlling the spatial patterns of SOC,soil aggregation,and aggregate stability in this region.The resistant and stable C fractions rather than labile C fractions are the major determinant of SOC stocks and aggregate stability.     

Degradation induces changes in the soil C:N:P stoichiometry of alpine steppe on the Tibetan Plateau

Due to the Tibetan Plateau's unique high altitude and low temperature climate conditions,the region's alpine steppe ecosystem is highly fragile and is suffering from severe degradation under the stress of increasing population,overgrazing,and climate change.The soil stoichiometry,a crucial part of ecological stoichiometry,provides a fundamental approach for understanding ecosystem processes by examining the relative proportions and balance of the three elements.Understanding the impact of degradation on the soil stoichiometry is vital for conservation and management in the alpine steppe on the Tibetan Plateau.This study aims to examine the response of soil stoichiometry to degradation and explore the underlying biotic and abiotic mechanisms in the alpine steppe.We conducted a field survey in a sequent degraded alpine steppe with seven levels inNorthern Tibet.The plant species,aboveground biomass,and physical and chemical soil properties such as the moisture content,temperature,pH,compactness,total carbon(C),total nitrogen(N),and total phosphorus(P)were measured and recorded.The results showed that the contents of soil C/N,C/P,and N/P consistently decreased along intensifying degradation gradients.Using regression analysis and a structural equation model(SEM),we found that the C/N,C/P,and N/P ratios were positively affected by the soil compactness,soil moisture content and species richness of graminoids but negatively affected by soil pH and the proportion of aboveground biomass of forbs.The soil temperature had a negative effect on the C/N ratio but showed positive effect on the C/P and N/P ratios.The current study shows that degradation-induced changes in abiotic and biotic conditions such as soil warming and drying,which accelerated the soil organic carbon mineralization,as well as the increase in the proportion of forbs,whichwere difficult to decompose and input less organic carbon into soil,resulted in the decreases in soil C/N,C/P,and N/P contents to a great extent.Our results provide a sound basis for sustainable conservation and management of the alpine steppe.     

Effects of grazing exclusion on plant productivity and soil carbon,nitrogen storage in alpine meadows in northern Tibet,China

Grazing exclusion is widely adopted in restoring degraded alpine grasslands on the Qinghai-Tibetan Plateau. However, its effectiveness remains poorly understood. In this study, we investigated the effects of grazing exclusion on plant productivity, species diversity and soil organic carbon （SOC） and soil total nitrogen （STN） storage along a transect spanning from east to west of alpine meadows in northern Tibet, China. After six years of grazing exclusion, plant cover, aboveground biomass （AGB）, belowground biomass （BGB）, SOC and STN were increased, but species diversity indices declined. The enhancement of AGB and SOC caused by grazing exclusion was correlated positively with mean annual precipitation （MAP）. Grazing exclusion led to remarkable biomass increase of sedge species, especially Kobresia pygmaea, whereas decrease of biomass in forbs and no obvious change in grass, leguminous and noxious species. Root biomass was concentrated in the near surface layer （10 cm） after grazing exclusion. The effects of grazing exclusion on SOC storage were confined to shallow soil layer in sites with lower MAP. It is indicated that grazing exclusion is an effective measure to increase forage production and enhance soil carbon sequestration in the studied region. The effect is more efficient in sites with higher precipitation. However, the results revealed a tradeoff between vegetation restoration and ecological biodiversity. Therefore, carbon pools recover more quickly than plant biodiversity in the alpine meadows. We suggest that grazing exclusion should be combined with other measures to reconcile grassland restoration and biodiversitv conservation.     

Trait complementarity between fine roots of Stipa purpurea and their associated arbuscular mycorrhizal fungi along a precipitation gradient in Tibetan alpine steppe

Development of fine roots and formation of symbiosis with arbuscular mycorrhizal(AM) fungi represent two strategies for plants to acquire nutrient and water from soil. Here, we elucidated how fine root development and symbolized mycorrhizal fungi with Stipa purpurea responded to the precipitation change in Tibetan alpine steppe ecosystem across a precipitation gradient from 50 mm to 400 mm. As precipitation increased, the proportion of thinner fine roots(diameter 0.4 mm) in total roots increased significantly; while the mycorrhizal colonization percentage, either associated with thinner or thicker roots, decreased. This phenomenon indicated that fine root development and symbolized mycorrhizal fungi are likely alternative, and plant preferred to develop fine root rather than build a symbiotic relationship with mycorrhizal fungi in more benign niches with higher precipitation. Also, root diameter was negatively correlated with specific root length(SRL), but positively correlated with AM fungal colonization percentage, indicating thicker-root species rely more on mycorrhizal fungi in alpine steppe. The complementarity between fine root and mycorrhizal fungi of S. purpurea is mediated by precipitation in Tibetan alpine steppe.     

Degradation significantly decreased the ecosystem multifunctionality of three alpine grasslands: evidences from a large-scale survey on the Qinghai-Tibetan Plateau

Owing to the joint effects of ecosystem fragility,anthropogenic disturbance and climate change,alpine grasslands(alpine meadow,alpine steppe and alpine desert)have experienced serious degradation during the past several decades.Grasslands degradation has severely affected the delivery of ecosystem multifunctionality(EMF)and services,and then threatens the livelihood of local herdsmen and ecological security of China.However,we still lack comprehensive insights about the effects of degradation and climatic factors on EMF of alpine grasslands,especially for alpine desert ecosystem.Therefore,we applied a large-scale field investigation to answer this question.Our results suggested grassland degradation significantly decreased the belowground ecosystem multifunctionality(BEMF)and EMF of alpine grasslands and aboveground ecosystem multifunctionality(AEMF)of alpine meadow,while did not reduce the AEMF of alpine steppe and desert.Except for the insignificant difference between degraded steppe and degraded desert in AEMF,the alpine meadow showed the highest AEMF,BEMF and EMF,alpine steppe ranked the second and alpine desert was the lowest.AEMF,BEMF and EMF of health alpine grasslands were strongly affected by mean annual precipitation(MAP)(19%-51%)and mean annual temperature(MAT)(9%-36%),while those of degraded meadow and degraded desert were not impacted by precipitation and temperature.AEMF and BEMF showed a synergistic relationship in healthy alpine grasslands(12%-28%),but not in degraded grasslands.Our findings emphasized the urgency of implementing the feasible ecological restoration project to mitigate the negative influences of grassland degradation on EMF of alpine ecosystems.     

Response of biomass spatial pattern of alpine vegetation to climate change in permafrost region of the Qinghai-Tibet Plateau,China

Alpine ecosystems in permafrost region are extremely sensitive to climate changes.To determine spatial pattern variations in alpine meadow and alpine steppe biomass dynamics in the permafrost region of the Qinghai-Tibet Plateau,China,calibrated with historical datasets of above-ground biomass production within the permafrost region's two main ecosystems,an ecosystem-biomass model was developed by employing empirical spatialdistribution models of the study region's precipitation,air temperature and soil temperature.This model was then successfully used to simulate the spatio-temporal variations in annual alpine ecosystem biomass production under climate change.For a 0.44°C decade-1 rise in air temperature,the model predicted that the biomasses of alpine meadow and alpine steppe remained roughly the same if annual precipitation increased by 8 mm per decade-1,but the biomasses were decreased by 2.7% and 2.4%,respectively if precipitation was constant.For a 2.2°C decade-1 rise in air temperature coupled with a 12 mm decade-1 rise in precipitation,the model predicted that the biomass of alpine meadow was unchanged or slightly increased,while that of alpine steppe was increased by 5.2%.However,in the absence of any rise in precipitation,the model predicted 6.8% and 4.6% declines in alpine meadow and alpine steppe biomasses,respectively.The response of alpine steppe biomass to the rising air temperatures and precipitation was significantly lesser and greater,respectively than that of alpine meadow biomass.A better understanding of the difference in alpine ecosystem biomass production under climate change is greatly significant with respect to the influence of climate change on the carbon and water cycles in the permafrost regions of the Qinghai-Tibet Plateau.     

Biomass allocation patterns of alpine grassland species and functional groups along a precipitation gradient on the Northern Tibetan Plateau

Variations in the fractions of biomass allocated to functional components are widely considered as plant responses to resource availability for grassland plants. Observations indicated shoots isometrically relates to roots at the community level but allometrically at the species level in Tibetan alpine grasslands. These differences may result from the specific complementarity of functional groups between functional components, such as leaf, root, stem and reproductive organ. To test the component complementary responses to regional moisture variation, we conducted a multi-site transect survey to measure plant individual size and component biomass fractions of common species belonging to the functional groups： forbs, grasses, legumes and sedges on the Northern Tibetan Plateau in peak growing season in 2010. Along the mean annual precipitation （MAP） gradient, we sampled 7o species, in which 2o are in alpine meadows, 20 in alpine steppes, 15 in alpine desert-steppes and 15 in alpine deserts, respectively. Our results showed that the size of alpine plants is small with individual biomass mostly lower than 1.0 g. Plants keep relative conservative component individual responses moisture functional fractions across alpine grasslands at the level. However, the complementary between functional components to variations specifically differ among groups. These results indicate that functional group diversity may be an effective tool for scaling biomass allocation patterns from individual up to community level. Therefore, it is necessary andvaluable to perform intensive and systematic studies on identification and differentiation the influences of compositional changes in functional groups on ecosystem primary services and processes.     

Soil organic carbon and nutrients along an alpine grassland transect across Northern Tibet

Soil carbon and nutrient contents and their importance in advancing our understanding of biogeochemical cycling in terrestrial ecosystem, has motivated ecologists to find their spatial patterns in various geographical area. Few studies have focused on changes in the physical and chemical properties of soils at high altitudes. Our aim was to identify the spatial distribution of soil physical and chemical properties in cold and arid climatic region. We also tried to explore relationship between soil organic carbon (SOC) and total nitrogen (TN), total phosphorus (TP), available nitrogen (AN), available phosphorus (AP), soil particle size distribution (PSD). Samples were collected at 44 sites along a 300 km transect across the alpine grassland of northern Tibet. The study results showed that grassland type was the main factor influencing SOC, TN and TP distribution along the Gangdise Mountain-Shenzha-Shuanghu Transect. SOC, TN and TP contents were significantly higher in alpine meadow than alpine steppe ecosystems. SOC, TN, TP and AN contents in two soil layers (0-15 cm and 15-30 cm) showed no significant differences, while AP content in top soil (0-15 cm) was significantly higher than that in sub-top soil (15-30 cm). SOC content was correlated positively with TN and TP content (r = 0.901 and 0.510, respectively). No correlations were detected for clay content and fractal dimension of particle size distribution (D). Our study results indicated the effects of vegetation on soil C, N and P seem to be more important than that of rocks itself along latitude gradient on the northern Tibetan Plateau. However, we did not found similar impacts of vegetation on soil properties in depth. Inaddition, this study also provided an interesting contribution to the global data pool on soil carbon stocks.     

Contrasting changes in above- and below-ground biomass allocation across treeline ecotones in southeast Tibet

Under conditions of a warmer climate, the advance of the alpine treeline into alpine tundra has implications for carbon dynamics in mountain ecosystems. However, the above- and below-ground live biomass allocations among different vegetation types within the treeline ecotones are not well investigated. To determine the altitudinal patterns of above-/below-ground carbon allocation, we measured the root biomass and estimated the above-ground biomass (AGB) in a subalpine forest, treeline forest, alpine shrub, and alpine grassland along two elevational transects towards the alpine tundra in southeast Tibet. The AGB strongly declined with increasing elevation, which was associated with a decrease in the leaf area index and a consequent reduction in carbon gain. The fine root biomass (FRB) increased significantly more in the alpine shrub and grassland than in the treeline forest, whereas the coarse root biomass changed little with increasing altitudes, which led to a stable below-ground biomass (BGB) value across altitudes. Warm and infertile soil conditions might explain the large amount of FRB in alpine shrub and grassland. Consequently, the root to shoot biomass ratio increased sharply with altitude, which suggested a remarkable shift of biomass allocation to root systems near the alpine tundra. Our findings demonstrate contrasting changes in AGB and BGB allocations across treeline ecotones, which should be considered when estimating carbon dynamics with shifting treelines.     

IMPACT OF ROAD CONSTRUCTION ON VEGETATION ALONGSIDE QINGHAI-XIZANG HIGHWAY AND RAILWAY

Based on the field investigation in August 2001 and August 2002, digital China Vegetation Map in 2001 and Qinghai-Xizang(Tibet) Plateau Vegetation Regionalization Map in 1996, vegetation characteristics along two sides of Qinghai-Xizang highway and railway are studied in this paper. Meanwhile, the impact of Qinghai-Xizang highway and railway constructions on the vegetation types are analyzed using ARCVIEW. ARC/1NFO and PATCH ANALYSIS. It was found that: 1) Qinghai-Xizang highway and railway span 9 latitudes, 12 longitudes and 6 physical geographic regions (East Qinghai and Qilian mountain steppe region, Qaidam mountain desert region,South Qinghai-Xizang alpine meadow steppe region. Qiangtang alpine steppe region, Golog-Nagqu alpine shrubmeadow region and South Xizang mountain shrub steppe region); 2) the construction of Qinghai-Xizang highway and railway destroyed natural vegetation and landscape, especially in 50m-wide buffer regions along both sides of the roads, it was estimated that the net primary productivity deceased by about 30 504.62t/a and the gross biomass deceased by 432 919.25-1 436 104.3t. The losing primary productivity accounted for 5.70% of the annual primary productivity within lkm-wide buffer regions (535 005.07-535 740.11t/a), and only 0.80%-0.89% of that within 10km-wide buffer regions (3 408 950.45-3 810 480.92t/a). The losing gross biomass was about 9.47%-17.06% of the gross biomass within lkm-wide buffer regions (7 502 971.85-25 488 342.71t), and only 1.47%-2.94% of that within 10km-wide buffer regions (43 615 065.35-164 150 665.37t).     

Effect of mixture sowing on biomass allocation in the artificially-planted pastures,Southeastern Tibetan

Artificial planting is an important measure to promote the restoration of degraded grassland and protect the ecological environment. The objectives of the current study were to investigate the allocation pattern between aboveground biomass(AGB) and belowground biomass(BGB) in different seeding types of artificially-planted pastures. We explored the variation in biomass and the relationship between above-and belowground biomass in four artificiallyplanted pastures with one species from Elymus nutans Griseb(EN, perennial), Elymus sibiricus Linn(ES, perennial), Medicago sativa Linn(MS, perennial), and Avena sativa Linn(AS, annual) and in six artificially-planted communities with mixtures of two species by seeding ratio 1:1 from the abovementioned grasses(EN + AS, MS + AS, EN + ES, MS + EN, MS + ES, AS + ES) in 2015 and 2016. The results showed that E. nutans is the most productive species with the highest biomass production among the single crops. MS + ES was the most productive group in 2015, while the group with the highest biomass production changed to AS + ES in 2016. AGB was positively correlated to BGB in the surface soil layer in the first year, but positively related to BGB in the subsoil layer in the second year. In the early stageof artificial grassland succession, plants allocated more biomass to aboveground parts, with a root to shoot(R/S) ratio of 1.98. The slope of the log-log relationship between AGB and BGB was 1.07 in 2016, which is consistent with the isometric theory. Different sowing patterns strongly affected the accumulation and allocation of biomass in artificiallyplanted grassland, E. sibiricus was the suitable plant in the alpine regions, which will be conducive to understanding vegetation restoration and plant interactions in the future.     

Aboveground Biomass and Water Storage Allocation in Alpine Willow Shrubs in the Qilian Mountains in China

The aboveground biomass allocation and water relations in alpine shrubs can provide useful information on analyzing their ecological and hydrological functions in alpine regions. The objectives of this study were to compare the aboveground biomass allocation, water storage ratio and distribution between foliage/woody components,and to investigate factors affecting aboveground biomass allocation and water storage ratio in alpine willow shrubs in the Qilian Mountains, China. Three experimental sites were selected along distance gradients from the riverside in the Hulu watershed in the Qilian Mountains. The foliage, woody component biomass, and water allocation of Salix cupularis Rehd.and Salix oritrepha Schneid. shrubs were measured using the selective destructive method. The results indicated that the foliage component had higher relative water and biomass storage than the woody component in the upper part of the crown in individual shrubs. However, the woody component was the major biomass and water storage component in the whole shrub level for S. cupularis and S.oritrepha. Moreover, the foliage/woody component biomass ratio decreased from the top to the basal level of shrubs. The relative water storage allocation was significantly affected by species types, but was not affected by sites and interaction between species and sites. Meanwhile, relative water storage was affectedby sites as well as by interaction between sites and species type.     

Aboveground biomass of the alpine shrub ecosystems in Three-River Source Region of the Tibetan Plateau

Though aboveground biomass (AGB) has an important contribution to the global carbon cycle, the information about storage and climatic effects of AGB is scare in Three-River Source Region (TRSR) shrub ecosystems. This study investigated AGB storage and its climatic controls in the TRSR alpine shrub ecosystems using data collected from 23 sites on the Tibetan Plateau from 2011 to 2013. We estimated the AGB storage (both shrub layer biomass and grass layer biomass) in the alpine shrubs as 37.49 Tg, with an average density of 1447.31 g m^-2. Biomass was primarily accumulated in the shrub layer, which accounted for 92% of AGB, while the grass layer accounted for only 8%. AGB significantly increased with the mean annual temperature (P < 0.05). The effects of the mean annual precipitation on AGB were not significant. These results suggest that temperature, rather than precipitation, has significantly effects on of aboveground vegetation growth in the TRSR alpine shrub ecosystems. The actual and potential increase in AGB density was different due to global warming varies among different regions of the TRSR. We conclude that long-term monitoring of dynamic changes is necessary to improve the accuracy estimations of potential AGB carbon sequestration across the TRSR alpine shrub ecosystems.     

Climatic and geographic factors affect ecosystem multifunctionality through biodiversity in the Tibetan alpine grasslands

Ecosystem multifunctionality(EMF), the simultaneous provision of multiple ecosystem functions, is often affected by biodiversity and environmental factors. We know little about how the interactions between biodiversity and environmental factors affect EMF. In this case study, a structural equation model was used to clarify climatic and geographic pathways that affect EMF by varying biodiversity in the Tibetan alpine grasslands. In addition to services related to carbon, nitrogen, and water cycling, forage supply, which is related to plantproductivity and palatability, was included in the EMF index. The results showed that 72% of the variation in EMF could be explained by biodiversity and other environmental factors. The ratio of palatable richness to all species richness explained 8.3% of the EMF variation. We found that air temperature, elevation, and latitude all affected EMF, but in different ways. Air temperature and elevation impacted the aboveground parts of the ecosystem, which included plant height, aboveground biomass, richness of palatable species, and ratio of palatable richness to all species richness. Latitude affected EMF by varying both aboveground and belowground parts of the ecosystem, which included palatable speciesrichness and belowground biomass. Our results indicated that there are still uncertainties in the biodiversity–EMF relationships related to the variable components of EMF, and climatic and geographic factors. Clarification of pathways that affect EMF using structural equation modeling techniques could elucidate the mechanisms by which environmental changes affect EMF.     

Dynamics of above- and below-ground biomass and C, N, P accumulation in the alpine steppe of Northern Tibet

The temporal dynamics of the biomass, as well as the carbon (C), nitrogen (N), phosphorus (P) concentrations and accumulation contents, in aboveand below-ground vegetation components were determined in the alpine steppe vegetation of Northern Tibet during the growing season of 2010. The highest levels of total biomass (311.68 g m⁻²), total C (115.95 g m⁻²), total N (2.60 g m⁻²), and total P (0.90 g m⁻²) accumulation contents were obtained in August in 2010. Further, biomass and nutrient stocks in the below-ground components were higher than those of the above-ground components. The dominant species viz., Stipa purpurea and Carex moorcrofti had lower biomass and C, N, P accumulations than the companion species which including Oxytropis. spp., Artemisia capillaris Thunb., Aster tataricus L., and so on.     

The Response of Vegetation Biomass to Soil Properties along Degradation Gradients of Alpine Meadow at Zoige Plateau

Alpine grassland of the Tibetan Plateau has undergone severe degradation, even desertification. However, several questions remain to be answered, especially the response mechanisms of vegetation biomass to soil properties. In this study, an experiment on degradation gradients was conducted in an alpine meadow at the Zoige Plateau in 2017. Both vegetation characteristics and soil properties were observed during the peak season of plant growth. The classification and regression tree model(CART) and structural equation modelling(SEM) were applied to screen the main factors that govern the vegetation dynamics and explore the interaction of these screened factors. Both aboveground biomass(AGB) and belowground biomass(BGB) experienced a remarkable decrease along the degradation gradients. All soil properties experienced significant variations along the degradation gradients at the 0.05 significance level. Soil physical and chemical properties explained 54.78% of the variation in vegetation biomass along the degradation gradients. AGB was mainly influenced by soil water content(SWC), soil bulk density(SBD), soil organic carbon(SOC), soil total nitrogen(STN), and pH. Soil available nitrogen(SAN), SOC and p H, had significant influence on BGB. Most soil properties had positive effects on AGB and BGB, while SBD and p H had a slightly negative effect on AGB and BGB. The correlations of SWC with AGB and BGB were relatively less significant than those of other soil properties. Our results highlighted that the soil properties played important roles in regulating vegetation dynamics along the degradation gradients and that SWC is not the main factor limiting plant growth in the humid Zoige region. Our results can provide guidance for the restoration and improvement of degraded alpine grasslands on the Tibetan Plateau.     

Degradation stage effects on vegetation and soil properties interactions in alpine steppe

In recent decades,overgrazing and the warming and drying climate have resulted in significant degradation of alpine grasslands in the source region of the Yellow River.However,research into the relationships between vegetation and soil properties has mainly focused on an overall degradation stage,and few studies have investigated which soil properties can impact vegetation change at different stages of degradation.Vegetation and soil properties were observed in the field and measured in the laboratory for different predefined stages of degradation for alpine grassland in Maduo County in the source region of the Yellow River.Results show that:1)the aboveground and belowground biomass,soil organic carbon,total nitrogen,nitrate,and ammonia content did not decrease significantly from the intact to moderate degradation stage,but decreased significantly at severe and very severe stages of degradation;2)dominant plant species shifted from gramineaes and sedges to forbs;3)the species richness and Pielou evenness indices decreased significantly at the very severe degradation stage,the Shannon-Wiener diversity index increased at the slight and moderate degradation,but decreased at the severe and very severe degradation stages;4)soil bulk density was the strongest soil driver for changes in the plant biomass and community diversity at the intact,slight and moderate degradation stages,whereas soil organic carbon and nitrate nitrogen content were the main driving factors for changes in plant biomass and diversity at the severe and very severe degradation stages.Our results suggest that there may be different interactions between soil properties and plants before and after moderate stages of degradation.     

Adaptations of the floral characteristics and biomass allocation patterns of Gentiana hexaphylla to the altitudinal gradient of the eastern Qinghai-Tibet Plateau

This study addressed the floral component traits and biomass allocation patterns of Gentiana hexaphylla as well as the relationships of these parameters along an elevation gradient(approximately 3700 m, 3800 m, 3900 m, and 4000 m) on the eastern Qinghai-Tibet Plateau. The plant height, floral characteristics, and biomass allocation of G. hexaphylla were measured at different altitudes after field sampling, sorting, and drying. Plant height was significantly greater at 3700 m than that at other elevations. Flower length was significantly greater at 4000 m than that at other elevations, whereas the flower length at low elevations showed no significant differences. Corolla diameter increased with altitude, although the difference was not significant between 3800 m and 3900 m. Variations in biomass accumulation, including the aboveground, photosynthetic organ, flower and belowground biomasses, showed non-linear responses to changes in altitude. The aboveground and photosynthetic organ biomasses reached their lowest values at 4000 m, whereas the belowground and flower biomassreached minimum values at 3700 m. The sexual reproductive allocation of G. hexaphylla also increased with altitude, with a maximum observed at 4000 m. These results suggest that external environmental factors and altitudinal gradients as well as the biomass accumulation and allocation of G. hexaphylla play crucial roles in plant traits and significantly affect the ability of this species to adapt to harsh environments. The decreased number of flowers observed at higher altitudes may indicate a compensatory response for the lack of pollinators at high elevations, which is also suggested by the deformed flower shapes at high altitudes. In addition, the individual plant biomass(i.e., plant size) had significantly effect on flower length and corolla diameter. Based on the organ biomass results, the optimal altitude for G. hexaphylla in the eastern Qinghai-Tibet Plateau is 3800 m, where the plant exhibits minimum propagule biomass and asexual reproductive allocation.     

Adaptations of the floral characteristics and biomass allocation patterns of <Emphasis Type="Italic">Gentiana hexaphylla</Emphasis> to the altitudinal gradient of the eastern Qinghai-Tibet Plateau

This study addressed the floral component traits and biomass allocation patterns of Gentiana hexaphylla as well as the relationships of these parameters along an elevation gradient (approximately 3700 m, 3800 m, 3900 m, and 4000 m) on the eastern Qinghai-Tibet Plateau. The plant height, floral characteristics, and biomass allocation of G. hexaphylla were measured at different altitudes after field sampling, sorting, and drying. Plant height was significantly greater at 3700 m than that at other elevations. Flower length was significantly greater at 4000 m than that at other elevations, whereas the flower length at low elevations showed no significant differences. Corolla diameter increased with altitude, although the difference was not significant between 3800 m and 3900 m. Variations in biomass accumulation, including the aboveground, photosynthetic organ, flower and belowground biomasses, showed non-linear responses to changes in altitude. The aboveground and photosynthetic organ biomasses reached their lowest values at 4000 m, whereas the belowground and flower biomass reached minimum values at 3700 m. The sexual reproductive allocation of G. hexaphylla also increased with altitude, with a maximum observed at 4000 m. These results suggest that external environmental factors and altitudinal gradients as well as the biomass accumulation and allocation of G. hexaphylla play crucial roles in plant traits and significantly affect the ability of this species to adapt to harsh environments. The decreased number of flowers observed at higher altitudes may indicate a compensatory response for the lack of pollinators at high elevations, which is also suggested by the deformed flower shapes at high altitudes. In addition, the individual plant biomass (i.e., plant size) had significantly effect on flower length and corolla diameter. Based on the organ biomass results, the optimal altitude for G. hexaphylla in the eastern Qinghai-Tibet Plateau is 3800 m, where the plant exhibits minimum propagule biomass and asexual reproductive allocation.     

Impacts of warming on root biomass allocation in alpine steppe on the north Tibetan Plateau

Biomass is an important component of global carbon cycling and is vulnerable to climate change. Previous studies have mainly focused on the responses of aboveground biomass and phenology to warming, while studies of root architecture and of root biomass allocation between coarse and fine roots have been scarcely reported in grassland ecosystems. We conducted an open-top-chamber warming experiment to investigate the effect of potential warming on root biomass and root allocation in alpine steppe on the north Tibetan Plateau. The results showed that Stipa purpurea had significantly higher total root length, root surface area and tips than Carex moocroftii. However,there were no differences in total root volume, mean diameter and forks for the two species. Warming significantly increased total root biomass(27.60%), root biomass at 0–10 cm depth(27.84%) and coarse root biomass(diameter 0.20 mm, 57.68%) in the growing season(August). However, warming had no significant influence on root biomass in the non-growing season(April). Root biomass showed clear seasonalvariations: total root biomass, root biomass at 0–10 cm depth and coarse root biomass significantly increased in the growing season. The increase in total root biomass was due to the enhancement of root biomass at 0–10 cm depth, to which the increase of coarse root biomass made a great contribution. This research is of significance for understanding biomass allocation, carbon cycling and biological adaptability in alpine grassland ecosystems under future climate change.