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

Energy exchange between the atmosphere and a meadow ecosystem on the Qinghai-Tibetan Plateau

To reveal the potential contribution of grassland ecosystems to climate change, we examined the energy exchange over an alpine Kobresia meadow on the northeastern Qinghai-Tibetan Plateau. The annual pattern of energy exchange showed a clear distinction between periods of frozen soil with the daily mean soil temperature at 5 cm (T-s5 &LE; 0 &DEG; C) and non-frozen soil (T-s5 > 0 &DEG; C). More than 80% of net radiation was converted to sensible heat (H) during the frozen soil period, but H varied considerably with the change in vegetation during the non-frozen soil period. Three different sub-periods were further distinguished for the later period: (1) the pre-growth period with Bowen ratio (β) > 1 was characterized by a high β of 3.0 in average and the rapid increase of net radiation associated with the increases of H, latent heat (LE) and soil heat; (2) during the Growth period when β &LE; 1, the LE was high but H fluxes was low with β changing between 0.3 and 0.4; (3) the post-growth period with average β of 3.6 when H increased again and reached a second maximum around early October. The seasonal pattern suggests that the phenology of the vegetation and the soil water content were the major factors affecting the energy partitioning in the alpine meadow ecosystem. © 2005 Elsevier B.V. All rights reserved.

Genetic variation within and among populations of Rhodiola alsia (Crassulaceae) native to the Tibetan Plateau as detected by ISSR markers

Genetic variation of 10 Rhodiola alsia ( Crassulaceae) populations from the Qinghai - Tibet Plateau of China was investigated using intersimple sequence repeat (ISSR) markers. R. alsia is an endemic species of the Qinghai - Tibet Plateau. Of the 100 primers screened, 13 were highly polymorphic. Using these primers, 140 discernible DNA fragments were generated with 112 (80%) being polymorphic, indicating pronounced genetic variation at the species level. Also there were high levels of polymorphism at the population level with the percentage of polymorphic bands (PPB) ranging from 63.4 to 88.6%. Analysis of molecular variance (AMOVA) showed that the genetic variation was mainly found among populations (70.3%) and variance within populations was 29.7%. The main factors responsible for the high level of differentiation among populations are probably the isolation from other populations and clonal propagation of this species. Occasional sexual reproduction might occur in order to maintain high levels of variation within populations. Environmental conditions could also influence population genetic structure as they occur in severe habitats. The strong genetic differentiation among populations in our study indicates that the conservation of genetic variability in R. alsia requires maintenance of as many populations as possible.

RAPD profiling in detecting genetic variation in endemic Coelonema (Brassicaceae) of Qinghai-Tibet Plateau of China

Random amplified polymorphic DNA ( RAPD) markers were used to measure genetic diversity of Coelonema draboides ( Brassicaceae), a genus endemic to the Qilian Mountains of the Qinghai-Tibet Plateau. We sampled 90 individuals in 30 populations of Coelonema draboides from Datong and Huzhu counties of Qinghai Province in P. R. China. A total of 186 amplified bands were scored from the 14 RAPD primers, with a mean of 13.3 amplified bands per primer, and 87% ( 161 bands) polymorphic bands (PPB) was found. Analysis of molecular variance (AMOVA) shows that a large proportion of genetic variation (84.2%) resides among individuals within populations, while only 15.8% resides among populations. The species shows higher genetic diversity between individuals than other endemic and endangered plants. The RAPDs provide a useful tool for assessing genetic diversity of rare, endemic species and for resolving relationships among populations. The results show that the genetic diversity of this species is high, possibly allowing it to adapt more easily to environmental variations. The main factor responsible for the high level of differentiation within populations and the low level of diversity among populations is probably the outcrossing and long-lived nature of this species. Some long-distance dispersal, even among far separated populations, is also a crucial determinant for the pattern of genetic variation in the species. This distributive pattern of genetic variation of C. draboides populations provides important baseline data for conservation and collection strategies for the species. It is suggested that only populations in different habitats should be studied and protected, not all populations, so as to retain as much genetic diversity as possible.

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.

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.

Experimental warming causes large and rapid species loss, dampened by simulated grazing, on the Tibetan Plateau

We investigated the independent and combined effects of experimental warming and grazing on plant species diversity on the north-eastern Tibetan Plateau, a region highly vulnerable to ongoing climate and land use changes. Experimental warming caused a 26-36% decrease in species richness, a response that was generally dampened by experimental grazing. Higher species losses occurred at the drier sites where N was less available. Moreover, we observed an indirect effect of climate change on species richness as mediated by plant-plant interactions. Heat stress and warming-induced litter accumulation are potential explanations for the species' responses to experimental warming. This is the first reported experimental evidence that climate warming could cause dramatic declines in plant species diversity in high elevation ecosystems over short time frames and supports model predictions of species losses with anthropogenic climate change.

Seasonal patterns of gross primary production and ecosystem respiration in an alpine meadow ecosystem on the Qinghai-Tibetan Plateau

We measured the net ecosystem CO2 exchange (NEE) in an alpine meadow ecosystem (latitude 37degrees29'-45'N, longitude 101degrees12'-23'E, 3250 m above sea level) on the Qinghai-Tibetan Plateau throughout 2002 by the eddy covariance method to examine the carbon dynamics and budget on this unique plateau. Diurnal changes in gross primary production (GPP) and ecosystem respiration (R-e) showed that an afternoon increase of NEE was highly associated with an increase of R-e. Seasonal changes in GPP corresponded well to changes in the leaf area index and daily photosynthetic photon flux density. The ratio of GPP/R-e was high and reached about 2.0 during the peak growing season, which indicates that mainly autotrophic respiration controlled the carbon dynamics of the ecosystem. Seasonal changes in mean GPP and R-e showed compensatory behavior as reported for temperate and Mediterranean ecosystems, but those of GPP(max) and R-emax were poorly synchronized. The alpine ecosystem exhibited lower GPP (575 g C m(-2) y(-1)) than, but net ecosystem production (78.5 g C m(-2) y(-1)) similar to, that of subalpine forest ecosystems. The results suggest that the alpine meadow behaved as a CO2 sink during the 1-year measurement period but apparently sequestered a rather small amount of C in comparison with similar alpine ecosystems.

Carbon dioxide exchange between the atmosphere and an alpine meadow ecosystem on the Qinghai-Tibetan Plateau, China

We used the eddy covariance method to measure the M exchange between the atmosphere and an alpine meadow ecosystem (37degrees29-45'N, 101degrees12-23'E, 3250m a.s.l.) on the Qinghai-Tibetan Plateau, China in the 2001 and 2002 growing seasons. The maximum rates Of CO2 uptake and release derived from the diurnal course Of CO2 flux (FCO2) were -10.8 and 4.4 mumol m(-2) s(-1), respectively, indicating a relatively high net carbon sequestration potential as compared to subalpine coniferous forest at similar elevation and latitude. The largest daily CO2 uptake was 3.9 g cm(-2) per day on 7 July 2002, which is less than half of those reported for lowland grassland and forest at similar latitudes. The daily CO2 uptake during the measurement period indicated that the alpine ecosystem might behave as a sink of atmospheric M during the growing season if the carbon lost due to grazing is not significant. The daytime CO2 uptake was linearly correlated with the daily photosynthetic photon flux density each month. The nighttime averaged F-CO2 showed a positive exponential correlation with the soil temperature, but apparently negative correlation with the soil water content. (C) 2004 Elsevier B.V. All rights reserved.

Methane emissions from different vegetation zones in a Qinghai-Tibetan Plateau wetland

We measured methane (CH4) emissions in the Luanhaizi wetland, a typical alpine wetland on the Qinghai-Tibetan Plateau, China, during the plant growth season (early July to mid-September) in 2002. Our aim was to quantify the spatial and temporal variation of CH4 flux and to elucidate key factors in this variation. Static chamber measurements of CH4 flux were made in four vegetation zones along a gradient of water depth. There were three emergent-plant zones (Hippuris-dominated; Scirpus-dominated; and Carex-dominated) and one submerged-plant zone (Potamogeton-dominated). The smallest CH4 flux (seasonal mean = 33.1 mg CH4 m(-2) d(-1)) was, observed in the Potamogeton-dominated zone, which occupied about 74% of the total area of the wetland. The greatest CH4 flux (seasonal mean = 214 mg CH4 m(-2) d(-1)) was observed in the Hippuris-dominated zone, in the second-deepest water area. CH4 flux from three zones (excluding the Carex-dominated zone) showed a marked diurnal change and decreased dramatically under dark conditions. Light intensity had a major influence on the temporal variation in CH4 flux, at least in three of the zones. Methane fluxes from all zones increased during the growing season with increasing aboveground biomass. CH4 flux from the Scirpus-dominated zone was significantly lower than in the other emergent-plant zones despite the large biomass, because the root and rhizome intake ports for CH4 transport in the dominant species were distributed in shallower and more oxidative soil than occupied in the other zones. Spatial and temporal variation in CH4 flux from the alpine wetland was determined by the vegetation zone. Among the dominant species in each zone, there were variations in the density and biomass of shoots, gas-transport system, and root-rhizome architecture. The CH4 flux from a typical alpine wetland on the Qinghai-Tibetan Plateau was as high as those of other boreal and alpine wetlands. (C) 2004 Elsevier Ltd. All rights reserved.

The impact of plateau zokor Myospalax fontanierii burrows on alpine meadow vegetation on the Qinghai-Xizang (Tibetan) plateau

The impact of burrows constructed by plateau zokors Myospalax fontanierii (Milne-Edwards, 1867) on alpine meadow vegetation on the Qinghai-Xizang (Tibetan) plateau was investigated. Plant samples taken from quadrats directly over active zokor burrows, back-filled burrows, adjacent burrow controls, and random sites from a field, in which no burrows or mounds occurred were compared. The biomass of plants (below- and above-ground) directly over shallow active burrows was significantly lower than on control plots. This reduction in biomass was not significantly different than that between deep active burrows and control plots. There were no significant differences between above- and below-ground plant biomass on areas perpendicular to active burrows when compared to random sites. Back-filling soil in burrows could promote the growth of above-ground monocotyledonous plants. However, the burrowing activities of zokors had a negative effect on biomass of dicotyledonous plants.

Uniformity of karyotypes in Ligularia (Asteraceae : Senecioneae), a highly diversified genus of the eastern Qinghai-Tibet Plateau highlands and adjacent areas

Ligularia, a highly diversified genus in the eastern Qinghai-Tibet Plateau and adjacent areas, was chosen as a suitable subject in which to study speciation patterns in this 'hot spot' area at the chromosomal level. Chromosome numbers and karyotypes were studied in 23 populations of 14 species, most of which are endemic to this area. The basic number x = 29 was confirmed for all species. Ligularia virgaurea was found to have diploid and triploid cytotypes, 2n = 58 and 87. Other species are only diploid, with 2n = 58. The karyotypes of all populations within any species, and all species spanning most sections and covering most of the morphological range in Ligularia, are very similar to each other, belonging to type 2A according to Stebbin's classification. This karyotype was also found in its close allies, e.g. Cremanthodium, Ligulariopsis, Parasenecio, and Sinacalia. Aneuploid reduction of chromosome number from 2n = 60 to 58 and karyotypic variation was found in Ligularia and its allies. Such a chromosomal pattern with few polyploids infers that variation of karyotype structure at the diploid level seems to be the predominant feature of chromosomal evolution in this group and sympatric speciation via hybridization and polyploidization has played a minor role in its species diversity. (C) 2004 The Linnean Society of London

Cloning of hypoxia-inducible factor 1 alpha cDNA from a high hypoxia tolerant mammal - plateau pika (Ochotona curzoniae)

Hypoxia-inducible factor I is a transcription factor composed of HIF-1alpha and HIF-1beta. It plays an important role in the signal transduction of cell response to hypoxia. Plateau pika (Ochotona curzoniae) is a high hypoxia-tolerant and cold adaptation species livin only at 3000-5000m above sea level on the Qinghai-Tibet Plateau. In this study, HIF-1alpha cDNA of plateau pika was cloned and its expression in various tissues was studied. The results indicated that plateau pika HIF-1alpha cDNA was highly identical to those of the human (82%), bovine (89%), mouse (82%), and Norway rat (77%). The deduced amino acid sequence (822 bp) showed 90%, 92%, 86%, and 86% identities with those of the human, bovine, house mouse, and Norway rat, respectively. Northern blot analyses detected two isoforms named pLHIF-1alpha and pSHIF-1alpha. The HIF-1alpha mRNA was highly expressed in the brain and kidney, and much less in the heart, lung, liver, muscle, and spleen, which was quite different from the expression pattern of mouse mRNA. Meanwhile, a new variant of plateau pika HIF-1alpha mRNA was identified by RT-PCR and characterized. The deduced protein, composed of 536 amino acids, lacks a part of the oxygen-dependent degradation domain (ODD), both transactivation domains (TADs), and the nuclear localization signal motif (NLS). Our results suggest that HIF-1alpha may play an important role in the pika's adaptation to hypoxia, especially in brain and kidney, and pika HIF-1alpha function pattern may be different from that of mouse HIF-1alpha. Further-more, for the high ratio of HIF-1alpha homology among the animals, the HIF-1alpha gene may be a good phylogenetic performer in recovering the true phylogenetic relationships among taxa. (C) 2004 Elsevier Inc. All rights reserved.

Grazing intensity alters soil respiration in an alpine meadow on the Tibetan plateau

Grazing intensity may alter the soil respiration rate in grassland ecosystems. The objectives of our study were to (1) determine the influence of grazing intensity on temporal variations in soil respiration of an alpine meadow on the northeastern Tibetan Plateau; and (2) characterise, the temperature response of soil respiration under different grazing intensities. Diurnal and seasonal soil respiration rates were measured for two alpine meadow sites with different grazing intensities. The light grazing (LG) meadow site had a grazing intensity of 2.55 sheep ha(-1), while the grazing intensity of the heavy grazing (HG) meadow site, 5.35 sheep ha(-1), was approximately twice that of the LG site. Soil respiration measurements - showed that CO2 efflux was almost twice as great at the LG site as at the HG site during the growing season, but the diurnal and seasonal patterns of soil respiration rate were similar for the two sites. Both exhibited the highest annual soil respiration rate in mid-August and the lowest in January. Soil respiration rate was highly dependent on soil temperature. The Q(10) value for annual soil respiration was lower for the HG site (2.75) than for the LG site (3.22). Estimates of net ecosystem CO2 exchange from monthly measurements of biomass and soil respiration revealed that during the period from May 1998 to April 1999, the LG site released 2040 g CO2 m(-2) y(-1) to the atmosphere, which was about one third more than the 1530g CO2 m(-2) y(-1) released at the HG site. The results suggest that (1) grazing intensity alters not only soil respiration rate, but also the temperature dependence of soil CO2 efflux; and (2) soil temperature is the major environmental factor controlling the temporal variation of soil respiration rate in the alpine meadow ecosystem. (C) 2003 Elsevier Ltd. All fights reserved.