A stable carbon isotopic approach for understanding the CO2 flux at the Haibei Alpine Meadow Ecosystem - A simple model

Although respiration of organisms and biomass as well as fossil fuel burning industrial production are identified as the major sources, the CO2 flux is still unclear due to the lack of proper measurements. A mass-balance approach that exploits differences in the carbon isotopic signature (delta(13)C) of CO2 Sources and sinks was introduced and may provide a means of reducing uncertainties in the atmospheric budget. delta(13)C measurements of atmospheric CO2 yielded an average of - 10.3 parts per thousand relative to the Peedee Belemnite standard; soil and plants had a narrow range from -25.09 parts per thousand to -26.51 parts per thousand and averaged at -25.80 parts per thousand. Based on the fact of steady fractionation and enrichment during respiration of mitochondria, we obtained the emission Of CO2 of 35.451 mol m(-2) a(-1) and CO2 flux of 0.2149 mu mol m(-2) s(-)1. The positive CO2 flux indicated the Haibei Alpine Meadow Ecosystem a source rather than a sink. The mass-balance model can be applied for other ecosystem even global carbon cycles because it neglects the complicated process of carbon metabolism, however just focuses on stable carbon isotopic compositions in any of compartments of carbon sources and sinks. (C) 2005 Elsevier B.V. All rights reserved.

Modeling trophic positions of the alpine meadow ecosystem combining stable carbon and nitrogen isotope ratios

Stable carbon and nitrogen isotope ratios of single tissues or whole bodies were analyzed to establish trophic positions of main consumers living at the alpine meadow ecosystem in the Tibetan Plateau. The results demonstrated that delta C-13 and delta N-15 values of vertebrates showed great variations and ranged from -26.83 to -22.51 parts per thousand and from 2.33 to 8.44 parts per thousand, respectively. Plateau pika, root vole, plateau hare, infants of rodents and hatchlings of passerine bird species had the lowest delta C-13 and delta N-15 values. delta C-13 and delta N-15 values of omnivorous and insectivorous birds and amphibians showed intermediate. Carnivorous species, steppe polecat and Upland buzzard, and omnivorous Robin accentor and White wagtail possessed extremely higher VC and delta N-15 values. Omnivorous birds captured in earlier year had significantly less negative delta C-13 and greater delta N-15 values than those captured later. Based on steady angular enrichment between trophic levels, an "alpha and vector model" combing delta C-13 and delta N-15 values was introduced to reveal trophic positions, the results indicated that Tibetan sheep, Tibetan yak, plateau pika, root vole, plateau hare, infants of small rodents showed the lowest trophic positions (TP 1.81-2.38). While omnivorous and insectivorous birds, their hatchlings and amphibians showed intermediate trophic positions (TP 2.06-2.89), carnivorous species steppe polecat and Upland buzzard, migrant birds possessed extremely higher trophic positions (TP 2.89-3.05). The isotopic investigation of organisms and the introduced "alpha and vector model" successfully demonstrated the same trophic positions and diet prediction of consumers as nitrogen enrichment model at the alpine meadow ecosystem. Besides of this information, the "alpha and vector model" can also be incorporated into multiple isotope signatures to infer trophic relationships. This angular enrichment model has the potential to address basic ecological questions, such as trophic structure, trophic dynamics, and energy flow in other terrestrial ecosystems of properly handled. (C) 2005 Elsevier B.V. All rights reserved.

The effect of land management on plant community composition, species diversity, and productivity of alpine Kobersia steppe meadow

Grassland degradation is widespread and severe on the Tibet Plateau. To explore management approaches for sustainable development of degraded and restored ecosystems, we studied the effect of land degradation on species composition, species diversity, and vegetation productivity, and examined the relative influence of various rehabilitation practices (two seeding treatments and a non-seeded natural recovery treatment) on community structure and vegetation productivity in early secondary succession. The results showed: (1) All sedge and grass species of the natural steppe meadow had disappeared from the severely degraded land. The above-ground and root biomass of severely degraded land were only 38 and 14.7%, respectively, of those of the control. So, the original ecosystem has been dramatically altered by land degradation on alpine steppe meadow. (2) Seeding measures may promote above-ground biomass, particularly grass biomass, and ground cover. Except for the grasses seeded, however, other grass and sedge species did not occur after seeding treatments in the sixth year of seeding. Establishment of grasses during natural recovery treatment progressed slowly compared with during seeding treatments. Many annual forbs invaded and established during the 6 years of natural recovery. In addition, there was greater diversity after natural recovery treatment than after seeding treatments. (3) The above-ground biomass after seeding treatment and natural recovery treatment were 114 and 55%, respectively, of that of the control. No significant differences in root biomass occurred among the natural recovery and seeded treatments. Root biomass after rehabilitation treatment was 23-31% that of the control.

Stable carbon isotope analyses show dominance of C3 grasses in the feed selected by small herbivores at the alpine meadow at the Tibetan Plateau

Muscle samples were collected from small herbivorous mammals (Ochotona curzoniae, Microtus oecnomus, Myospalax fontanierii and Lepus oiostolus) at the alpine meadow ecosystem at the Tibetan Plateau in order to address variability in stable carbon isotope composition. Stable carbon isotope values of muscles remain steady and show no significant variations (-25.72 to -27.04 parts per thousand) among the four small mammal species. Based on the mass balance theory of stable isotopes, it is proposed that small herbivorous mammals mainly (or totally) rely on C3 grasses as food supply, and there is few or no distribution of C4 grasses at the ecosystem. The results reflect our previous study on the isotope patterns of plant species. Thus, stable carbon isotope analysis of muscles provides a method to address dietary selection and dietary variability in herbivores. In addition, stable carbon isotopic analyses can be used to address changes in vegetation distributions in ecosystem and paleovegetaion and paleoclimate.

Floral closure in response to temperature and pollination in Gentiana straminea Maxim. (Gentianaceae), an alpine perennial in the Qinghai-Tibetan Plateau

Floral closure may be induced by pollination and various other factors, but is rarely studied comprehensively. Different kinds of floral closure should have various effects on reproductive fitness of plants. Two contrasting types of floral closure were observed in the flowers of Gentiana straminea Maxim. in the eastern Qinghai-Tibetan Plateau. The first type occurred prior to pollination during both gender phases, in response mainly to decreasing air temperatures. Flowers closed when decreasing temperatures approached 20 degrees C and subsequently began to reopen the following day during mid-morning when air temperatures warmed to approximately 13-15 degrees C. This kind of floral closure can protect pollen grains on either stamens or stigmas, increasing fitness of both male and female. Following pollination, permanent floral closure occurred, although there was a delay between the dates of pollination and permanent closure, during which flowers continued to show temporary closure in response to low temperature episodes. The time required for permanent, pollination-induced closure varied according to the age of the gender phase, including a prolonged time before closure if pollination occurred early in the female phase. The retaining of permanent closed flowers increased both approaching (to inflorescences) and visiting (to unpollinated flowers) frequencies of individual plants when with fewer open flowers and the persisting corolla is further beneficial for seed sets of these pollinated flowers. Thus, two separate types of floral closure, one in response to environmental cues and the other in response to the age of each gender stage, appeared to have a strong influence on reproductive fitness in this species. These results revealed a different adaptive strategy of alpine plants in the sexual reproduction assurance in addition to the well-known elevated floral longevity, dominant role of more effective pollinators and increased reproduction allocation in the arid habitats.

Effect of burrowing activity of root vole (Microtus oeconomus Pallas) on plant diversity in alpine meadow

Herbivory and burrowing activity of mammals may influence the species composition and diversity of plant communities. The effect of corridors and holes systems constructed by root vole (Microtus oeconomus Pallas) on the plant species diversity was studied in the habitat of high - mountain meadow (3250 in a.s.l in Qinghai-Tibet Plateau, China). By using grid method, these disturbances were studied on 16 plots (100 cm x 100 cm) distributed in 4 transects in studied area, in August 2000 and 2001. The disturbance intensity index, D, was calculated as the percent of the ground surface disturbed by voles in the study area. Plant species were identified and counted on the same plots. In total 46 plant species were identified - 39% of this number was considered as sensitive to the vole disturbances as their occurrence and/or abundance decreased along the disturbance intensity. Generally, a significantly negative correlation (r = - 0.911 P < 0.01) between vole aboveground disturbances and plant species diversity (H') was found. The results suggest that root vole ground disturbances, especially in the form of actively utilized holes and corridors have significantly negative influence on plant species diversity in high-mountain grassland habitat.

The effect of land management on carbon and nitrogen status in plants and soils of alpine meadows on the Tibetan plateau

Large-scale grassland rehabilitation has been carried out on the severely degraded lands of the Tibetan plateau. The grasslands created provide a useful model for evaluating the recovery of ecosystem properties. The purposes of this research were: (1) to examine the relative influence of various rehabilitation practices on carbon and nitrogen in plants and soils in early secondary succession; and (2) to evaluate the degree to which severely degraded grassland altered plant and soil properties relative to the non-disturbed native community. The results showed: (1) The aboveground tissue C and N content in the control were 105-97 g m(-2) and 3.356gm(-2), respectively. The aboveground tissue C content in the mixed seed treatment, the single seed treatment, the natural recovery treatment and the severely degraded treatment was 137 per cent, 98 per cent, 49 per cent and 38 per cent, respectively, of that in the control. The corresponding aboveground tissue N content was 109 per cent, 84 per cent, 60 per cent and 47 per cent, respectively, of that in the control. (2) Root C and N content in 0-20 cm depths of the control had an 2 2 average 1606 gm(-2) and 30-36 gm(-2) respectively. Root C and N content in the rehabilitation treatments were in the range of 26-36 per cent and 35-53 per cent, while those in the severely degraded treatment were only 17 per cent and 26 per cent of that in the control. (3) In the control the average soil C and N content at 0-20 cm was 11307 gm(-2) and 846 gm(-2), respectively. Soil C content in the uppermost 20 cm in the seeded treatments, the natural recovery treatment and the severely degraded treatment was 67 per cent, 73 per cent and 57 per cent, respectively, while soil N content in the uppermost 20cm was 72 per cent, 82 per cent and 79 per cent, respectively, of that in the control. The severely degraded land was a major C source. Restoring the severely degraded lands to perennial vegetation was an alternative approach to sequestering C in former degraded systems. N was a limiting factor in seeding grassland. It is necessary for sustainable utilization of seeding grassland to supply extra N fertilizer to the soil or to add legume species into the seed mix. Copyright (c) 2005 John Wiley & Sons, Ltd.

The potential importance of grazing to the fluxes of carbon dioxide and methane in an alpine wetland on the Qinghai-Tibetan Plateau

To assess the impact of livestock grazing on the emission of greenhouse gases from grazed wetlands, we examined biomass growth of plants, CO2 and CH4 fluxes under grazing and non-grazing conditions on the Qinghai-Tibetan Plateau wetland. After the grazing treatment for a period of about 3 months, net ecosystem CO2 uptake and aboveground biomass were significantly smaller, but ecosystem CH4 emissions were remarkably greater, under grazing conditions than under non-grazing conditions. Examination of the gas-transport system showed that the increased CH4 emissions resulted from mainly the increase of conductance in the gas-transport system of the grazed plants. The sum of global warming potential, which was estimated from the measured CO2 and CH4 fluxes, was 5.6- to 11.3-fold higher under grazing conditions than under non-grazing conditions. The results suggest that livestock grazing may increase the global warming potential of the alpine wetlands. (c) 2005 Elsevier Ltd. All rights reserved.

Effect of annual weeds on the growth of perennial grass mixtures in the alpine region of the Qinghai-Tibetan Plateau

In the alpine region of the Tibetan Plateau, five perennial grass cultivars, Bromus inermis (B), Elymus nutans (E), Clinelymus nutans (C), Agropyron cristatum (A), and Poa crymophila (P) were combined into nine communities with different compositions and ratios, B+C, E+A, B+E+A, E+B+C,C+E+A,B+E+C+A,B+C+A+P,B+E+A+P and E+C+A+P. Each combination was sown in six 10 X 10 m plots with three hand-weeded plots and three natural-growing plots in a completely randomised design in 1998. A field experiment studied the performance of these perennial grass combinations under the competitive interference of annual weeds in 3 consecutive years from 1998 to 2000. The results showed that annual weeds occupied more space and suppressed the growth of the grasses due to earlier germination and quicker growth in the establishment year, but this pattern changed in the second and third years. Leaf area indexes (LAIs) of grasses were greatly decreased by the competitive interference of weeds, and the negative effect of weeds on LAIs of grasses declined and stabilised in the second and third years. E+B+C, B+E+C+A, and B+E+A+P possessed relatively higher LAIs (P < 0.05) among all grass combinations and their LAIs were close to five when the competitive interference of weeds was removed. Grasses were competitively inferior to weeds in the establishment year, although their competitive ability (aggressivities) increased throughout the growing season. In the second and third years, grasses were competitively superior to weeds, and their competitive ability decreased from May until August and increased in September. Dry matter (DM) yields of grasses were reduced by 29.8-74.1% in the establishment year, 11.0-64.9% in the second year, and 16.0-55.8% in the third year by the competitive interference of weeds. B+E+C+A and B+E+A+P can produce around 14 t/ha of DM yields, significantly higher (P < 0.05) than the production of the other grass combinations in the second and third years after the competitive interference of weeds was removed. It was preliminarily concluded that removal of competitive interference of weeds increased the LAIs of all grass swards and improved the light interception of grasses, thus promoting the production of perennial grass pastures. The germination stage of the grasses in the establishment year was the critical period for weeding and suppression of weeds should occur at an early stage of plant growth. The grass combinations of B+E+C+A and B+E+A+P were productive and can be extensively established in the alpine regions of the Tibetan Plateau. Two or three growing seasons will be needed before determining success of establishment of grass mixtures under the alpine conditions of the Tibetan Plateau.

Effect of N fertilizer on the productivity and nutritive values of perennial grass mixtures in the alpine region of Qinghai-Tibetan Plateau, China

A study was conducted on grass mixtures that included smooth bromegrass (SB) + drooping wild ryegrass (DW), smooth bromegrass + Siberian wild ryegrass (SW) + crested wheatgrass (CW) and smooth bromegrass + Siberian wild ryegrass + drooping wild ryegrass + crested wheatgrass in the alpine region of Qinghai-Tibetan Plateau. The study was conducted from 1998 to 2000 to investigate the effects of N application rates and growing year on herbage dry matter (DM) yield and nutritive values. Herbage DM production increased linearly with N application rates. The effect of N application on DM yields was greater (P < 0.05) in the 2nd and 3rd production years than in the establishment year. Dry matter yields of SB + SW + CW and SB + SW + DW + CW can reach as high as 15 000 kg ha(-1) at 345 kg ha(-1) N rate in the 3rd growing year. With increased N application rates, crude protein (CP) contents and 48 h in sacco DM degradability of grasses increased (P < 0.05). No effect (P > 0.05) of N application was detected on organic matter (OM) and acid detergent fibre (ADF) concentration. It can be concluded that for increased biomass production in the alpine region of the Qinghai-Tibetan Plateau, a minimum of 345 kg N ha(-1) should be applied to grass stands in three split application of 115 kg N ha(-1), in early June, early July and late July

Local warming about 1.3 degrees C in alpine meadow has no effect on root vole (Microtus oeconomus L.) population during winter

The influence of air and soil warming on root vole (Microtus oeconomus L.) population was studied in winter period in top open chambers (OTC) (0.8-1.8 m(2) warmed by conical fiberglass material and situated in alpine meadow (3250 m) at Qinghai-Tibet Plateau, China. The OTCs were distributed on an area of 30 x 30 m of experimental warming site; another site of the same area was a control one. The root vole population was investigated on two pairs of sites in "low-grazing" and "high-grazing" (by sheep) parts of the meadow; mark-recapture method was used. The winter-season averaged air and soil temperature inside of the chambers were 1.3 degrees C higher than the temperature outside the chambers. The warming in the chambers had no statistically significant effect on root vole numbers, on average body mass of individual, and on average body mass of males and females. In conclusion, as small as 1.3 degrees C warming of soil and air introduced locally and on small (several m(2)) scale, in the alpine meadow habitat in winter period, has possibly no effect on root vole numbers and biomass.

Productivity and persistence of perennial grass mixtures under competition from annual weeds in the alpine region of the Qinghai-Tibetan Plateau

In the alpine region of the Qinghai-Tibetan Plateau four indigenous perennial grass species Bromus inermis (BI), Elymus sibiricus (ES), Elymus nutans (EN) and Agropyron cristatum (AC) were cultivated as three mixtures with different compositions and seeding rates, BI + EN, BI + ES + AC and BI + ES + EN + AC. From 1998 to 2001 there were three different weeding treatments: never weeded (CK); weeded on three occasions in the first year (1-y) and weeded on three occasions in both the first and second year (2-y) and their effect of grass combination and interactions on sward productivity and persistence was measured. Intense competitive interference by weedy annuals reduced dry matter (DM) yield of the swards. Grass combination significantly affected sward DM yields, leaf area index (LAI) and foliar canopy cover and also species composition DM and LAI, and species plant cover. Interaction between weeding treatments and grass combination was significant for sward DM yield, LAI and canopy cover, but not on species composition for DM, LAI or species plant cover. Grass mixture BI + ES + EN + AC gave the highest sward DM yield and LAI for both weeding and non-weeding treatments. Species ES and EN were competitively superior to the others. Annual weedy forbs must be controlled to obtain productive and stable mixtures of perennial grasses, and germination/emergence is the most important time for removal. Weeding three times (late May, late June and mid-July) in the establishment year is enough to maintain the production and persistence of perennial grass mixtures in the following growing seasons. Extra weeding three times in the second growing year makes only a slight improvement in productivity.

Nitrogen deposition and carbon sequestration in alpine meadows

Nitrogen deposition experiments were carried out in alpine meadow ecosystems in Qinghai-Xizang Plateau in China, in order to explore the contribution of nitrogen deposition to carbon sequestration in alpine meadows. Two methods were used in this respect. First, we used the allocation of N-15 tracer to soil and plant pools. Second, we used increased root biomass observed in the nitrogen-amended plots. Calculating enhanced carbon storage, we considered the net soil CO2 emissions exposed to nitrogen deposition in alpine meadows. Our results show that nitrogen deposition can enhance the net soil CO2 emissions, and thus offset part of carbon uptake by vegetation and soils. It means that we have to be cautious to draw a conclusion when we estimate the contribution of nitrogen deposition to carbon sequestration based on the partitioning of N-15 tracer in terrestrial ecosystems, in particular in N-limited ecosystems. Even if we assess the contribution of nitrogen deposition to carbon sequestration based on increased biomass exposed to nitrogen deposition in terrestrial ecosystems, likewise, we have to consider the effects of nitrogen deposition on the soil CO2 emissions.

Leaf orientation, incident sunlight, and photosynthesis in the alpine species Suassurea superba and Gentiana straminea on the Qinghai-Tibet Plateau

The extremely high level of solar radiation on the Qinghai-Tibet Plateau may induce photoinhibition and thus limit leaf carbon gain. To assess the effect of high light, we examined gas exchange and chlorophyll fluorescence for two species differing in light interception: the prostrate Saussurea superba and the erect-leaved Gentiana straminea. In controlled conditions with favorable water and temperature, neither species showed apparent photoinhibition in gas exchange measurements. In natural environment, however, their photosynthetic rate decreased remarkably at high light. Photosynthesis depression was aggravated under high leaf temperature or soil water stress. Relative stomatal limitation was much higher in S. superba than in G. straminea and it remarkably increased in the later species at midday when soil was dry. F-v/F-m as an indicator for photoinhibition was generally higher in S. superba than in the other species. F-v/F-m decreased significantly under high light at midday in both species, even when soil moisture was high. F-0 linearly elevated with the increment of leaf temperature in G. straminea, but remained almost constant in S. superba. Electron transport rate (ETR) increased with photosynthetically active photon flux density (PPFD) in S. superba, but declined when PPFD was high than about 1000 mumol m(-2) s(-1) in G. straminea. Compared to favorable environment, the estimated daily leaf carbon gain at PPFD above 800 mumol m(-2) s(-1) was reduced by 32% in S. superba and by 17% in G. straminea when soil was moist, and by 43% and 53%, respectively, when soil was dry. Our results suggest that the high radiation induces photoinhibition and significantly limits photosynthetic carbon gain, and the limitation may further increase at higher temperature and in dry soil.

Observation and modeling for terrestrial processes in alpine meadow

The water-heat transfer process between land and atmosphere in Haibei alpine meadow area has been systematically observed. A multi-layer coupling model for land-atmosphere interaction was presented with special attention paid to the moisture transfer in leaf stomata under unsaturated condition. A profound investigation on the physical process of turbulent transfer inside the vegetation has been performed with a revised formula of water absorption for root system. The present model facilitates the study of vertically distributed physical variables in detail. Numerical simulation was conducted according to the transfer process of Kinesia humility meadow in the area of Haibei Alpine Meadow Ecosystem Station, CAS. The calculated results agree well with observation.