Phylogeography of the Qinghai-Tibetan Plateau endemic Juniperus przewalskii (Cupressaceae) inferred from chloroplast DNA sequence variation

The vegetation of the northeast Qinghai-Tibetan Plateau is dominated by alpine meadow and desert-steppe with sparse forests scattered within it. To obtain a better understanding of the phylogeography of one constituent species of the forests in this region, we examined chloroplast trnT-trnF and trnS-trnG sequence variation within Juniperus przewalskii, a key endemic tree species. Sequence data were obtained from 392 trees in 20 populations covering the entire distribution range of the species. Six cpDNA haplotypes were identified. Significant population subdivision was detected (G(ST) = 0.772, N-ST = 0.834), suggesting low levels of recurrent gene flow among populations and significant phylogeographic structure (N-ST > G(ST), P < 0.05). Eight of the nine disjunct populations surveyed on the high-elevation northeast plateau were fixed for a single haplotype (A), while the remaining, more westerly population, contained the same haplotype at high frequency together with two low frequency haplotypes (C and F). In contrast, most populations that occurred at lower altitudes at the plateau edge were fixed or nearly fixed for one of two haplotypes, A or E. However, two plateau edge populations had haplotype compositions different from the rest. In one, four haplotypes (A, B, D and E) were present at approximately equivalent frequencies, which might reflect a larger refugium in the area of this population during the last glacial period. Phylogenetic analysis indicated that the most widely distributed haplotype A is not ancestral to other haplotypes. The contrasting phylogeographic structures of the haplotype-rich plateau edge area and the almost haplotype-uniform plateau platform region indicate that the plateau platform was recolonized by J. przewalskii during the most recent postglacial period. This is supported by the findings of a nested clade analysis, which inferred that postglacial range expansion from the plateau edge followed by recent fragmentation is largely responsible for the present-day spatial distribution of cpDNA haplotypes within the species.

Strong temperature dependence and no moss photosynthesis in winter CO2 flux for a Kobresia meadow on the Qinghai-Tibetan plateau

We examined the CO2 exchange of a Kobresia meadow ecosystem on the Qinghai-Tibetan plateau using a chamber system. CO2 efflux from the ecosystem was strongly dependence on soil surface temperature. The COZ efflux-temperature relationship was identical under both light and dark conditions, indicating that no photosynthesis could be detected under light conditions during the measurement period. The temperature sensitivity (Q(10)) of the COZ efflux showed a marked transition around -1.0 degrees C; Q(10) was 2.14 at soil surface temperatures above and equal to -1.0 degrees C but was 15.3 at temperatures below -1.0 degrees C. Our findings suggest that soil surface temperature was the major factor controlling winter COZ flux for the alpine meadow ecosystem and that freeze-thaw cycles at the soil surface layer play an important role in the temperature dependence of winter CO2 flux. (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

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.

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.

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.

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.

Short-term variation of CO2 flux in relation to environmental controls in an alpine meadow on the Qinghai-Tibetan Plateau

[1] The alpine meadow ecosystem on the Qinghai-Tibetan Plateau may play a significant role in the regional carbon cycle. To assess the CO2 flux and its relationship to environmental controls in the ecosystem, eddy covariance of CO2, H2O, and energy fluxes was measured with an open-path system in an alpine meadow on the plateau at an elevation of 3,250 m. Net ecosystem CO2 influx (Fc) averaged 8.8 g m(-2) day(-1) during the period from August 9 to 31, 2001, with a maximum of 15.9 g m(-2) day(-1) and a minimum of 2.3 g m(-2) day(-1). Daytime Fc averaged 16.7 g m(-2) day(-1) and ranged from 10.4 g m(-2) day(-1) to 21.7 g m(-2) day(-1) during the study period. For the same photosynthetic photon flux density (PPFD), gross CO2 uptake (Gc) was significantly higher on cloudy days than on clear days. However, mean daily Gc was higher on clear days than on cloudy days. With high PPFD, Fc decreased as air temperature increased from 10degreesC to 23degreesC. The greater the difference between daytime and nighttime air temperatures, the more the sink was strengthened. Daytime average water use efficiency of the ecosystem (WUEe) was 8.7 mg (CO2)(g H2O)(-1); WUEe values ranged from 5.8 to 15.3 mg (CO2)(g H2O)(-1). WUEe increased with the decrease in vapor pressure deficit. Daily albedo averaged 0.20, ranging from 0.19 to 0.22 during the study period, and was negatively correlated with daily Fc. Our measurements provided some of the first evidence on CO2 exchange for a temperate alpine meadow ecosystem on the Qinghai-Tibetan Plateau, which is necessary for assessing the carbon budget and carbon cycle processes for temperate grassland ecosystems.

New progress in the study of Paleozoic strata in the Xainza area, Tibet

The authors discovered and measured a relatively complete Paleozoic section (exposed continuously from the Lower Ordovician to Upper Permian) and collected plentiful fossils of many groups during geological investigations of the 1∶250000 Xainza County Sheet in the period of 2000-2002. It was the first time in Tibet that the representative element Tetragraptus (Paratetragraptus) approximatus of the graptolite zone at the lowermost part of the Early Ordovician Arenigian Stage and the typical elements Waagenophyllum indicum var. crassiseptatum and Liangshanophyllum streploseptatum Graptolithina of the Late Permian Wujiapingian coral fauna. Based on these, the Lower Ordovician Zakang Formation and Upper Permian Mujiu Co Formation were established. It not only improved the Paleozoic stratigraphic sequence in the area but also provided new information for the study of the Paleozoic tectonic movement and evolution of the Qinghai-Tibet Plateau.

Discovery of the Early Ordovician strata in Xianza County, Tibet and its significance

There is no any sediments contain fossils which formed earlier than Middle Ordovician in northern Tibet yet. It is very important whether have developed the Cambrian to Early Ordovician sedimentary strata. During geological mapping of Xainza County in 2000, Tetragraptus approximatus, the typical one of graptolite belts which cosmepolitically distributed in the Arenigian of Early Ordovician, was collected from the so-call Pre-Sinian Nianqingtanggula Group. So there is developped the Early Ordovician sediments and we have established Zakang Formation and Taduo Formation which perfected the Plaeozoic sequences in northern Tibet. According to these, we can realize tectonic control, palaeogeographrc pattern as well as palaeobiogeographic distribution of the Qinghai-Tibet Plateau in the early Paleozoic and also provided the materials to the cosmepolitical biostratigraphic contrast.