Seasonal changes in weight and body composition of yak grazing on alpine-meadow grassland in the Qinghai-Tibetan plateau of China

Forty-five male yaks (born April 2001) were studied to determine how seasonal changes on the Qinghai-Tibetan plateau affected BW and body composition. Thirty yaks were weighed monthly from birth to 26 mo of age to determine seasonal changes in BW. The remaining 15 yaks were allocated randomly to five groups (three yaks per group), designated for slaughter at 13, 15, 18, 22, and 25 mo to measure seasonal effects on body chemical composition. All yaks were grazed on the alpine-meadow grassland of the plateau without any supplementation. All BW and body composition data were calculated on an individual basis. Body weight and body composition data were both compared across seven growth periods spanning 2 yr and defined by season. From April (birth) to December 2001 of the first growing season, yak BW increased (P < 0.01); however, during the subsequent cold season (December 2001 to May 2002), BW decreased (P < 0.01). The second growing season ran from May 2002 (13 mo of age) to October 2002 (18 mo of age), and the second live weight-loss season ran from October 2002 until May 2003. The weight loss experienced by yaks during the first weight loss season was 25.64% of the total weight gain in the first growing season. The weight loss experienced by yaks during the second weight-loss season was 29.73% of the total weight gain in the second growing season. Energy retention in the second growing season was 291.07 MJ, 50.8% of which was consumed during the subsequent cold season. Energy accumulation in the summer (from May to July) and fall (from July to October) of the second growing season did not differ (5.01 and 6.30 MJ/kg of EBW gain, respectively; P = 0.63). The energy mobilized during the second winter (from October 2002 to February 2003) was 16.49 MJ/kg of EBW, and in the second spring (from February to May 2003), it was 9.06 MJ/kg of EBW. These data suggest that the decrease in grazing yak BW during the first cold season is much less than during the second cold season, and that the energy content per unit of BW mobilized is greater (P = 0.02) in winter than in spring. Results from this study demonstrate highly efficient compensatory growth in grazing yaks following the first weight loss period during the first cold season. This benefit could be exploited by herders to improve yak production. Yaks may have developed a type of self-protection mechanism to overcome the long cold seasons in the Qinghai-Tibetan plateau.

Origin of the Qinghai-Tibetan Plateau endemic Milula (Liliaceae): further insights from karyological comparisons with Allium

Milula, a monotypic genus endemic to the Qinghai-Tibetan Plateau, was found to be nested deeply within Allium by the molecular phylogeny despite the aberrant morphology. It remains unknown what had contributed to the rapid evolution of morphology and origin of this exceptional species. In contrast to a previous report of its karyotypes with 2n = 16 = 8M+8SM (2SAT), similar to most species of Allium, a rather different karyotype, 2n = 20 = 4M +10SM+6T (2SAT), was found in examined 31 individuals from 6 populations of M. spicata distributed in the central Tibet. Karyotypes of 7 Allium species occurring in the Qinghai-Tibetan Plateau were further reported. The basic number x = 8 was confirmed for all of them and their karyotypes consist mainly of metacentric and submetacentric chromosomes with rare subterminal and terminal chromosomes. The karyotype of M. spicata is distinctly different from that of most Allium species occurring in the plateau through a complete comparison of all available species in this region and adjacent areas. However, the same chromosome number and similar karyotypic structure were found in A. fasciculatum of Sect. Bromatorrhiza, indicating a possible close relationship between them. But this similarity is contradictory to the preliminary molecular phylogenetic analysis that Milula was closely related to A. cyathophorum of Sect. Bromatorrhiza with x=8, but the other species with x=10 and 11 in this section were clearly placed in the other clade. We therefore suggested that the paralleling evolution from x=8 to x=9, 10 and 11 with increasing asymmetry of karyotype possibly due to the chromosomal Robertsonian translocation might occur separately in the two recognized phylogenetic lineages of Allium. In addition to aneuploidy and following change of the chromosomal structures, the habitat isolation due to the recent uplift of the Qinghai-Tibetan Plateau and the Quaternary climatic oscillation, plays a greater role in origin of Milula and other endemic species (genera) with aberrant morphology from their progenitors.

Characterization of photosynthetic pathway of plant species growing in the eastern Tibetan plateau using stable carbon isotope composition

The photosynthetic pathway of plant species collected at Menyuan, Henan, and Maduo sites, east of Tibetan Plateau, China, during the growing season were studied using stable carbon isotopes in leaves. The 232 samples leaves analyzed belonged to 161 species, 30 families, and 94 genera. The delta(13)C values (from -24.6 to -29.2 %o) indicated that all the considered species had a photosynthetic C-3 pathway. The absence of plant species with C-4 photosynthetic pathway might be due to the extremely low air temperature characterizing the Tibetan Plateau. The average delta(13)C value was significantly (p < 0.05) different between annuals and perennials at the three considered study sites. Hence the longer-lived species had greater water-use efficiency (WUE) than shorter-lived species, that is, longer-lived species are better adapted to the extreme environmental conditions of the Tibetan Plateau.

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.

Radiation and diversification within the Ligularia-Cremanthodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau

The Ligularia-Cremanthodium-Parasenecio (L-C-P) complex of the Tussilagininae (Asteraceae: Senecioneae) contains more than 200 species that are endemic to the Qinghai-Tibetan Plateau in eastern Asia. These species are morphologically distinct; however, their relationships appear complex. A phylogenetic analysis of members of the complex and selected taxa, of the tribe Senecioneae was conducted using chloroplast (ndhF and trnL-F) and nuclear (ITS) sequences. Phylogenetic trees were constructed from individual and combined datasets of the three different sequences. All analyses suggested that Doronicum, a genus that has been included in the Tussilagininae, should be excluded from this subtribe and placed at the base of the tribe Senecioneae. In addition, the Tussilagininae should be broadly circumscribed to include the Tephroseridinae. Within the expanded Tussilagininae containing all 13 genera occurring in eastern Asia, Tussilago and NSPetasites diverged early as a separate lineage, while the remaining I I genera comprise an expanded L-C-P complex clade. We suggest that the L-C-P clade, which is largely unresolved, most likely originated as a consequence of an explosive radiation. The few monophyletic subclades identified in the L-C-P clade with robust support further suggest that some genera of Tussilagininae from eastern Asia require generic re-circumscriptions given the occurrence of subclades containing species of the same genus in different parts of the phylogentic tree due to homoplasy of important morphological characters used to delimit them. Molecular-clock analyses suggest that the explosive radiation of the L-C-P complex occurred mostly within the last 20 million years, which falls well within the period of recent major uplifts of the Qinghai-Tibetan Plateau between the early Miocene to the Pleistocene. It is proposed that significant increases in geological and ecological diversity that accompanied such uplifting, most likely promoted rapid and continuous allopatric speciation in small and isolated populations, and allowed fixation or acquisition of similar morphological characters within unrelated lineages. This phenomenon, possibly combined with interspecific diploid hybridization because of secondary sympatry during relatively stable stages between different uplifts, could be a major cause of high species diversity in the Qinghai-Tibetan Plateau and adjacent areas of eastern Asia. (c) 2005 Elsevier Inc. All rights reserved.

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.

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.

Reproductive ecology of the Qinghai-Tibet Plateau endemic Gentiana straminea (Gentianaceae), a hermaphrodite perennial characterized by herkogamy and dichogamy

The combined occurrence of both herkogamy and dichogamy in a hermaphrodite species has been considered to strongly favour outcrossing. In this study, we investigated in detail the reproductive ecology of Gentiana straminea Maxim. (Gentianaceae), a hermaphrodite perennial endemic to the Qinghai-Tibet Plateau. In a series of observations and experiments over four consecutive years, we examined whether the combination of dichogamy and herkogamy in individual flowers completely prevents geitonogamous pollen transfer in this species. The mode of floral development clearly indicates that autonomous self-pollination is completely avoided through herkogamy and dichogamy in individual flowers. This implication was confirmed by the breeding experiments, since no seed was produced when flowers were isolated. However, this gentian proved to be highly self-compatible when geitonogamous selfing was artificially induced. Many flowers opened simultaneously on individual plants, the ratio of male to female phase flowers was close to 2:1 in each inflorescence, at the full anthesis phase, and they were randomly distributed amongst the upper, middle and lower parts of each stem's inflorescence. On average, Bombus sushikini Skorikov, the most frequent visitor and only legitimate pollinator of G. straminea, visited nearly two flowers per inflorescence, and four flowers per plant. Among the pollinators' foraging bouts, the proportions of geitonogamous visits to inflorescences or flowers within an individual plant were 29% and 37%, respectively. Therefore, despite the strict dichogamous and herkogamous characteristics of the individual flowers, geitonogamous selfing might still prevail in G. straininea because of the size of its floral displays and the continuous visiting behavior of B. sushkini. (C) 2005 Elsevier SAS. All rights reserved.

Characterizing CO2 fluxes for growing and non-growing seasons in a shrub ecosystem on the Qinghai-Tibet Plateau

To assess carbon budget for shrub ecosystems on the Qinghai-Tibet Plateau, CO2 flux was measured with an open-path eddy covariance system for an alpine shrub ecosystem during growing and non-growing seasons. CO2 flux dynamics was distinct between the two seasons. During the growing season from May to September, the ecosystem exhibited net CO2 uptake from 08:00 to 19:00 (Beijing Standard Time), but net CO2 emission from 19:00 to 08:00. Maximum CO2 uptake appeared around 12:00 with values of 0.71, 1,19, 1.46 and 0.67 g CO2 m(-2) h(-1) for June, July, August and September, respectively. Diurnal fluctuation Of CO2 flux showed higher correlation with photosynthetic photon flux density than temperature. The maximum net CO2 influx occurred in August with a value of 247 g CO2 m(-2). The total CO2 uptake by the ecosystem was up to 583 g CO2 m(-2) for the growing season. During the non-growing season from January to April and from October to December, CO2 flux showed small fluctuation with the largest net CO2 efflux of 0.30 g CO2 m(-2) h(-1) in April. The diurnal CO2 flux was close to zero during most time of the day, but showed a small net CO2 eff lux from 11:00 to 18:00. Diurnal CO2 flux, is significantly correlated to diurnal temperature in the non-growing season. The maximum monthly net CO2 eff lux appeared in April, with a value of 105 g CO2 m(-2). The total net CO2 eff lux for the whole non-growing season was 356 g CO2 m(-2).

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.