Changes in individual plant traits and biomass allocation in alpine meadow with elevation variation on the Qinghai-Tibetan Plateau

Plant traits and individual plant biomass allocation of 57 perennial herbaceous species, belonging to three common functional groups (forbs, grasses and sedges) at subalpine (3700 m ASL), alpine (4300 m ASL) and subnival (>= 5000 m ASL) sites were examined to test the hypothesis that at high altitudes, plants reduce the proportion of aboveground parts and allocate more biomass to belowground parts, especially storage organs, as altitude increases, so as to geminate and resist environmental stress. However, results indicate that some divergence in biomass allocation exists among organs. With increasing altitude, the mean fractions of total biomass allocated to aboveground parts decreased. The mean fractions of total biomass allocation to storage organs at the subalpine site (7%+/- 2% S.E.) were distinct from those at the alpine (23%+/- 6%) and subnival (21%+/- 6%) sites, while the proportions of green leaves at all altitudes remained almost constant. At 4300 m and 5000 m, the mean fractions of flower stems decreased by 45% and 41%, respectively, while fine roots increased by 86% and 102%, respectively. Specific leaf areas and leaf areas of forbs and grasses deceased with rising elevation, while sedges showed opposite trends. For all three functional groups, leaf area ratio and leaf area root mass ratio decreased, while fine root biomass increased at higher altitudes. Biomass allocation patterns of alpine plants were characterized by a reduction in aboveground reproductive organs and enlargement of fine roots, while the proportion of leaves remained stable. It was beneficial for high altitude plants to compensate carbon gain and nutrient uptake under low temperature and limited nutrients by stabilizing biomass investment to photosynthetic structures and increasing the absorption surface area of fine roots. In contrast to forbs and grasses that had high mycorrhizal infection, sedges had higher single leaf area and more root fraction, especially fine roots.

Effects of temperature, glucose and inorganic nitrogen inputs on carbon mineralization in a Tibetan alpine meadow soil

High levels of available nitrogen (N) and carbon (C) have the potential to increase soil N and C mineralization We hypothesized that with an external labile C or N supply alpine meadow soil will have a significantly higher C mineralization potential and that temperature sensitivity of C mineralization will increase To test the hypotheses an incubation experiment was conducted with two doses of N or C supply at temperature of 5 15 and 25 C Results showed external N supply had no significant effect on CO2 emission However external C supply increased CO2 emission Temperature coefficient (Q(10)) ranged from 113 to 1 29 Significantly higher values were measured with C than with N addition and control treatment Temperature dependence of C mineralization was well-represented by exponential functions Under the control CO2 efflux rate was 425 g CO2-Cm-2 year(-1) comparable to the in situ measurement of 422 g CO2-Cm-2 year(-1) We demonstrated if N is disregarded microbial decomposition is primarily limited by lack of labile C It is predicted that labile C supply would further increase CO2 efflux from the alpine meadow soil (C) 2010 Elsevier Masson SAS All rights reserved

Vegetation Responses to A Long-term Grazing Intensity Experiment in Alpine Shrub Grassland on Qinghai-Tibet Plateau

在青藏高原中国科学院海北高寒草甸生态系统定位研究站对金露梅高寒灌丛草场植被开展了长期不同放 牧强度试验,分别在短期(4 年) 、中期(11 年) 和长期(18 年) 放牧阶段研究不同放牧干扰强度对草地植物物种多样 性、群落结构、地上生物量和草场质量的影响。研究表明,在不同放牧阶段,随着放牧强度增加植物群落的高度和 盖度都降低。在中期放牧干扰阶段,物种多样性指数和均匀度指数随着放牧强度增加呈现典型的单峰曲线模式; 在长期放牧干扰阶段,随着放牧强度增加,占优势地位的灌木和禾草被典型杂类草替代,其中的重度放牧干扰简化 了高寒灌丛植被群落结构,减少了地上现存生物量,特别是可食优良牧草生物量。植被对放牧的响应除了与放牧 强度和放牧时间阶段密切相关外,还与该地区水热条件的变化有一定的相关性。针对长期放牧干扰的反应特性可 将金露梅灌丛草场中植物划分为增加型、敏感型、忍耐型和无反应型4 种类型。除了丰富度指数、多样性指数和均 匀度指数外,其它一些特征参数并不支持著名的中度干扰假说。本研究发现,长期重度放牧促进了青藏高原高寒 草地退化,适度放牧有利于高寒灌丛草场的生物多样性保护和牧草利用;“取半留半”的放牧原则在青藏高原草场 放牧管理实践中值得推荐,它将有利于防止草场退化,提高牧草利用率和维持较高的生物多样性。

Surface energy partitioning in alpine swamp meadow in the Qinghai Tibetan Plateau

Based on surface energy flux data measured by eddy covariance methods from China Flux in alpine swamp meadow of the Qinghai Tibetan Plateau in 2005, the daily and seasonal dynamic of surface energy fluxes and their partitioning, as well as abiotic factors effects were analyzed. The results suggested that LE （Latent heat flux） was the largest consumer of the incoming energy. Rn （Net radiation flux） and LE showed clear seasonal variations in sharp hump and up to their maximums in August and July, respectively. H （Sensible heat flux） increased to its peak in August whereafter declined slowly. Precipitation could reduce the components of surface energy. As to Rn and LE, their correlations with abiotic factors were evident while it was not significant in H. Average EBR （Energy balance ratio） was 50.7 %, which was much larger in growing season than non-growing season.

Carbon Dioxide Exchange Between the Atmosphere and an Alpine Shrubland Meadow During the Growing Season on the Qinghai-Tibetan Plateau

In the present study, we used the eddy covariance method to measure CO2 exchange between the atmosphere and an alpine shrubland meadow ecosystem (37°36'N, 101°18'E; 3 250 m a.s.l.) on the Qinghai-Tibetan Plateau, China, during the growing season in 2003, from 20 April to 30 September. This meadow is dominated by formations of Potentilla fruticosa L. The soil is Mol-Cryic Cambisols. During the study period, the meadow was not grazed. The maximum rates of CO2 uptake and release derived from the diurnal course of CO2 flux were -9.38 and 5.02 μmol•m-2•s-1, respectively. The largest daily CO2 uptake was 1.7 g C•m-2•d-1 on 14 July, which is less than half that of an alpine Kobresia meadow ecosystem at similar latitudes. Daily CO2 uptake during the measurement period indicated that the alpine shrubland meadow ecosystem may behave as a sink of atmospheric CO2 during the growing season. The daytime CO2 uptake was correlated exponentially or linearly with the daily photo synthetic photon flux density each month. The daytime average water use efficiency of the ecosystem was 6.47 mg CO2/g H2O. The efficiency of the ecosystem increased with a decrease in vapor pressure deficit.

Stable Carbon Isotopic Composition in Soil Organic Carbon and C3/C4 Source Variations at the Haibei Alpine Meadow

Stable carbon isotopes of organic matter originated from different soil layers (0～5 cm, 5～15 cm, 15～25 cm, 25～35 cm, 35～50 cm, 50～65 cm) were investigated in the Haibei Alpine Meadow Ecosystem Research Station of the Chinese Academy of Sciences. The preliminary results indicated that δ13C values of soil organic matter increased with increased soil depth. δ13C of soil organic carbon in 0～5 cm layer showed the lowest value, -25.09‰; while 50～65 cm soil layer possessed the lowerδ13C value, -13.87‰. Based on mass balance model of stable isotopes, it was proposed that the percentage of C4 carbon source tend to increase with increased soil depth. The preliminary study indicated that alpine meadow might have undergone a successive process from C4-dominated community to C3-dominated one. However, changing δ13C values in atmospheric CO2 overtime and different processes of soil organic carbon formation (or eluviation) might somewhat contribute to increasing δ13C values. In this case, mass balance model would underestimate C3 community and overestimate C4 community.

Comparative study on CO2 emissions from different types of alpine meadows during grass exuberance period

Potentilla fruticosa scrub, Kobresia humilis meadow and Kobresia tibetica meadow are widely distributed on the Qinghai-Tibet Plateau. During the grass exuberance period from 3 July to 4September, based on close chamber-GC method, a study on CO2 emissions from different treatments was conducted in these meadows at Haibei research station, CAS. Results indicated that mean CO2emission rates from various treatments were 672.09+152.37 mgm-2h-1 for FC (grass treatment); 425.41+191.99 mgrn-2h-1 for FJ (grass exclusion treatment); 280.36+174.83 mgrn-2h-1 for FL (grass and roots exclusion treatment); 838.95+237.02 mgm-2h-1 for GG (scrub+grass treatment); 528.48+205.67 mgm-2h-1for GC (grass treatment); 268.97 ±99.72 mgm-2h-1 for GL (grass and roots exclusion treatment); and 659.20±94.83 mgm-2h-1 for LC (grass treatment), respectively (FC, FJ, FL, GG, GC, GL, LC were the Chinese abbreviation for various treatments). Furthermore, Kobresia humilis meadow, Potentilla fruticosa scrub meadow and Kobresia tibetica meadow differed greatly in average CO2 emission rate of soil-plant system, in the order of GG＞FC＞LC＞GC. Moreover, in Kobresia humilis meadow,heterotrophic and autotrophic respiration accounted for 42% and 58% of the total respiration of soil-plant system respectively, whereas, in Potentilla fruticosa scrub meadow, heterotrophic and autotrophic respiration accounted for 32% and 68% of total system respiration from G-G; 49% and 51%from GC. In addition, root respiration from Kobresia humilis meadow approximated 145 mgCO2m-2h-1,contributed 34% to soil respiration. During the experiment period, Kobresia humilis meadow and Potentilla fruticosa scrub meadow had a net carbon fixation of 111.11 grn-2 and 243.89 grn-2,respectively. Results also showed that soil temperature was the main factor which influenced CO2 emission from alpine meadow ecosystem, significant correlations were found between soil temperature at 5 cm depth and CO2 emission from GG, GC, FC and FJ treatments. In addition, soil moisture may be the inhibitory factor of CO2 emission from Kobresia tibetica meadow, and more detailed analyses should be done in further research.

No C_4 Plants Found at the Haibei Alpine Meadow Ecosystem Research Station in Qinghai, China: Evidence from Stable Carbon Isotope Studies

Using the measurement of stable carbon isotopes in leaves as a tool to investigate photosyn-thetic pathway of 102 plant species grown at an alpine meadow ecosystem, at the foot of the Qilian Mountain, Qinghai, China. The results indicate that the δ~(3)C values of plants have a narrow range from -28.24‰ to -24.84‰, which means that none of the species examined belongs to C_4 and crassulaceous acid metabolism (CAM) photosynthetic pathway and all of these species perform photosynthesis through the C_3 pathway. This is likely due to a long-term adaptation to environments at the alpine meadow ecosystem.

Effect of plateau zokors on vegetation characteristics and productivity of alpine meadow

This research was conducted on alpine meadow site at Menyuan county, Qinghai Province, People's Republic of China to determine the effects of native, subterranean rodent of Qinghai-Tibet grasslands, the plateau zokors (Myospalax baileyi), on seasonal above-and below-ground plant biomass, plant species diversity and productivity. Both total peaks of above-and below-ground biomass were the greatest (413.600 g/m~2 and 2297.502 g/m~2) in the patch no any plateau zokors colonized by plateau zokors over 10 years in August and October, respectively. Both above-and below-ground biomass were significantly increased in the patches where plateau zokors were removed or the burrow systems were abandoned for five years compared to the patches plateau zokors colonized over 10 years. However, both above-and below-ground biomass in abandoned patches were significantly lower than that in uncolonized patches. Monocotyledonous biomass was reduced greatly, but the non-palatable dicots were significantly increased in colonized patches. The palatable biomass of monocots and dicots were increased in abandoned patches. Total plant species diversity was the greatest in uncolonized patchesand least in abandoned patch. The total net primary production in colonized patches was reduced by 68.98% compared with uncolonized patches. Although the patches were without any plateau zokors disturbance for fives years, the total net primary production just reached 58.69% of the uncolonized patches. The above-ground net primary production in abandoned patches increased 28.74% and the below-ground increased 54.91% compared with the colonized patches. We suggest that plateau zokor-induced changes in plant above- and below-ground biomass and species diversity may lead to further alterations of nutrient cycling and trophic dynamics in this alpine meadow ecosystem.

Methane emissions by alpine plant communities in the Qinghai-Tibet Plateau

For the first time to our knowledge, we report here methane emissions by plant communities in alpine ecosystems in the Qinghai-Tibet Plateau. This has been achieved through long-term field observations from June 2003 to July 2006 using a closed chamber technique. Strong methane emission at the rate of 26.2 +/- 1.2 and 7.8 +/- 1.1 mu g CH4 m(-2) h(-1) was observed for a grass community in a Kobresia humilis meadow and a Potentilla fruticosa meadow, respectively. A shrub community in the Potentilla meadow consumed atmospheric methane at the rate of 5.8 +/- 1.3 mu g CH4 m(-2) h(-1) on a regional basis; plants from alpine meadows contribute at least 0.13 Tg CH4 yr(-1) in the Tibetan Plateau. This finding has important implications with regard to the regional methane budget and species-level difference should be considered when assessing methane emissions by plants.

Nitrous oxide emissions from two alpine meadows in the Qinghai-Tibetan Plateau

Nitrous oxide (N2O) emission was measured in a Kobresia humilis meadow and a Potentilla fruticosa meadow in the Qinghai-Tibet Plateau from June 2003 to July 2006. Five treatments were setup in the two alpine meadows. Two bare soil treatments were setup in the K. humilis meadow (BSK) and in the P. fruticosa meadow (BSP) by removing the above- and belowground plant biomass. Three plant community treatments were setup with one in the K. humilis meadow (herbaceous community in the K. humilis meadow-HCK) and two in the P. fruticosa meadow (herbaceous community in the P. fruticosa meadow-HCP, and shrub community in the P. fruticosa meadow-SCP). Nitrous oxide emission from BSP was estimated to be 38.1 +/- 3.6 mu g m(-2) h(-1), significantly higher than from BSK (30.2 +/- 2.8 mu g m(-2) h(-1)) during the whole experiment period. Rates from the two herbaceous blocks (HCK and HCP) were close to 39.5 mu g m(-2) stop h(-1) during the whole experimental period whereas shrub community (SCP) showed significant high emission rates of N2O. Annual rate of N2O emission was estimated to be 356.7 +/- 8.3 and 295.0 +/- 11.6 mg m(-2) year(-1) from the alpine P. fruticosa meadow and from the alpine K. humilis meadow, respectively. These results suggest that alpine meadows in the Qinghai-Tibetan Plateau are an important source of N2O, contributing an average of 0.3 Tg N2O year(-1). We concluded that N2O emission will decrease, due to a predicted vegetation shift from shrubs to grasses imposed by overgrazing.

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers' residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers' residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m(2) along a transect and classified the contents by species. We observed 15.5-19.7 species per 0.01 m(2), which is high richness per 0.01 m(2) on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.

Response of Polygonum viviparum species and community level to long-term livestock grazing in alpine shrub meadow in Qinghai-Tibet Plateau

Grazing by domestic herbivores is generally recognized as a major ecological factor and an important evolutionary force in grasslands. Grazing has both extensive and profound effects on individual plants and communities. We investigated the response patterns of Polygonum viviparum species and the species diversity of an alpine shrub meadow in response to long-term livestock grazing by a field manipulative experiment controlling livestock numbers on the Qinghai-Tibet Plateau in China. Here, we hypothesize that within a range of grazing pressure, grazing can alter relative allocation to different plant parts without changing total biomass for some plant species if there is life history trade-offs between plant traits. The same type of communities exposed to different grazing pressures may only alter relative species' abundances or species composition and not vary species diversity because plant species differ in resistant capability to herbivory. The results show that plant height and biomass of different organs differed among grazing treatments but total biomass remained constant. Biomass allocation and absolute investments to both reproduction and growth decreased and to belowground storage increased with increased grazing pressure, indicating the increasing in storage function was attained at a cost of reducing reproduction of bulbils and represented an optimal allocation and an adaptive response of the species to long-term aboveground damage. Moreover, our results showed multiform response types for either species groups or single species along the gradient of grazing intensity. Heavy grazing caused a 13.2% increase in species richness. There was difference in species composition of about 18%-20% among grazing treatment. Shannon-Wiener (H') diversity index and species evenness (E) index did not differ among grazing treatments. These results support our hypothesis.

Feeding behaviour of yaks on spring, transitional, summer and winter pasture in the alpine region of the Qinghai-Tibetan plateau

Of the present estimated world population of 14.2 million yaks, approximately 13.3 million occur within Chinese territories (Food and Agriculture Organization of the United Nations, 2003). Although there is an extensive bibliography covering the species, few studies have been conducted in the area of foraging behaviour. The present study was conducted at pasture during the spring, transitional, summer and winter seasons to determine the daily temporal patterns of grazing and ruminating behaviour by yaks. During each study period, two 24 h recordings were undertaken with each of six mature dairy yaks. One study period was conducted on each of the transitional, summer and winter pastures, whereas, due to the considerable changes occurring in the morphology of the spring pasture, three separate studies were completed during March, April and May. During the second of these studies (April), the effect of level of concentrate supplementation on grazing and ruminating behaviour was also examined. Behaviour recordings were made using solid-state behaviour recorders. Short-term intake rates (IR, g min(-1)) were calculated by weighing yaks before and after approximately 1 h of grazing, retaining the faeces and urine excreted and applying a correction for insensible weight loss. Yaks spent less time grazing during the dry season (the early period on the spring pasture) compared with the later green swards (the later period on the spring pasture, the transitional pasture and the summer pasture) (P < 0.05). When the forage quality improved, but there was still insufficient mass (the later period on the spring pasture), the yaks extended their grazing time at the expense of other activities. During the early periods on the spring pasture, the short-term IR by yaks was up to 53 g DM min(-1), significantly higher than at other times (P < 0.05). The level of concentrate offered had little or no effect on grazing or ruminating time. The total eating time of the yaks offered 0.5 or 1.0 kg concentrate was 2.9 and 4.5 h day(-1) respectively, significantly lower than unsupplemented yaks (6.8 h) (P < 0.05). In general, yaks can regulate their foraging behaviour according to the changes of sward conditions in order to achieve optimal grazing strategies. (C) 2007 Published by Elsevier B.V.

Potential refugium on the Qinghai-Tibet Plateau revealed by the chloroplast DNA phylogeography of the alpine species Metagentiana striata (Gentianaceae)

Metagentiana striata is an alpine annual herbaceous plant endemic to the east of the Qinghai-Tibet (Q-T) Plateau and adjacent areas. The phylogeography of M. striata was studied by sequencing the chloroplast DNA (cpDNA) trnS-trnG intergenic spacer. Ten haplotypes were identified from an investigation of 232 individuals of M. striata from 14 populations covering the entire geographical range of this species. The level of differentiation amongst populations was very high (G(ST) = 0.746; N-ST = 0.774) and a significant phylogeographical structure was observed (P < 0.05). An analysis of molecular variance found a high variation amongst populations (76%), with F-ST = 0.762 (highly significant, P < 0.001), indicating that little gene flow occurred amongst the different regions; this was explained by the isolation of populations by high mountains along the Q-T Plateau and adjacent areas (N-m = 0.156). Only one ancestral haplotype (A) was common and widespread throughout the distributional range of M. striata. The populations of the Hengduan Mountains region of the south-eastern Q-T Plateau showed high diversity and uniqueness of haplotypes. It is suggested that this region was the potential refugium of M. striata during the Quaternary glaciation, and that interglacial and postglacial range expansion occurred from this refugium. This scenario was in good agreement with the results of nested clade analysis, which inferred that the current spatial distribution of cpDNA haplotypes and populations resulted from range expansion, together with past allopatric fragmentation events. (c) 2008 The Linnean Society of London.