Belowground net primary productivity and biomass allocation of a grassland in Inner Mongolia is affected by grazing intensity

The root system of permanent grasslands is of outstanding importance for resource acquisition. Particularly under semi-arid conditions, the acquisition of water and nutrients is highly variable during the vegetation growth period and between years. Additionally, grazing is repeatedly disturbing the functional equilibrium between the root system and the transpiring leaf canopy. However, very few data is available considering grazing effects on belowground net primary productivity (BNPP) and root-shoot dry mass allocation in natural grassland systems. We hypothesise that grazing significantly reduces BNPP due to carbon reallocation to shoot growth. Root biomass and BNPP were estimated by soil coring in 2004, 2005 and 2006 and from ingrowth cores in 2005 and 2006 at one site which has been protected from grazing since 1979 (UG79), at one winter grazing (WG), and one heavily grazed (HG) site. BNPP was estimated from the summation of significant increments of total and live root biomass and from accumulated root biomass of ingrowth cores. Belowground biomass varied from 1,490–2,670 g m^?2 and was significantly lower under heavy grazing than at site UG79. Root turnover varied from 0.23 to 0.33 year^?1 and was not significantly different between sites. Heavy grazing significantly decreased live root biomass and BNPP compared to site UG79. Taking BNPP estimates from live root biomass dynamics and ingrowth cores as the most reliable values, the portion of dry mass allocated belowground relative to total net primary productivity (BNPP/NPP) varied between 0.50–0.66 and was reduced under heavy grazing in 2005, but not in 2006. The positive correlation between cumulative root length density of ingrowth cores and leaf dry matter suggests that the ingrowth core method is suitable for studying BNPP in this semi-arid steppe system. Grazing effects on BNPP and BNPP/NPP should be considered in regional carbon models and estimates of belowground nutrient cycling.     

Nitrogen, phosphorus and potassium nutritional status of semiarid steppe grassland in Inner Mongolia

In grazed semiarid steppe ecosystems, much attention has been paid to aspects of growth limitation by water. So far, potential limitation of primary production by plant nutrients was rarely considered. This knowledge is essential for identification of sustainable land-use practices in these large and important ecosystems on the background of over-exploitation and climate change. In the present study plant nutrient concentrations and ratios were investigated with factorial additions of water and N fertilizer at two sites with contrasting soil nutrient availability. Combined analysis of nutrient concentrations, contents, biomass production, and plant N:P ratios consistently confirmed primary growth limitation by water and a strong N limitation when sufficient amounts of water were supplied. P limitation only occurred at the site with low P availability when in addition to the natural supply, water and N fertilizer were given. According to reported thresholds of N:K and K:P ratios, K was not limiting in any plot. The observed nutritional patterns in the plant community were related to the dynamics of species composition and their specific nutrient status. Stipa grandis had the highest N:P ratio whereas Artemisia frigida showed lowest N:P. These nutrient characteristics were related to growth strategies of dominant species. Accordingly, the relative biomass contribution of S. grandis and A. frigida strongly affected the nutrient status of the plant community. Plant N:P ratios indicate the relative limitation by N or P in the semiarid grasslands under sufficient water supply, but other methods of nutritional diagnosis should be used when plant N:P ratios remain below critical values.     

Effects of N and water supply on water use-efficiency of a semiarid grassland in Inner Mongolia

Productivity of semiarid grasslands is primarily limited by seasonal rainfall amount and becomes increasingly limited by nutrient availability under wet conditions. Interactive effects of water and N availability on grassland productivity and parameters related to water use were studied on a grassland site in Inner Mongolia, China, in a 2-factorial experiment with two levels of water (rainfed: 158 mm; irrigated: 839 (N0) and 972 (N1) mm) and N supply (0 or 180 kg N ha^?1). RUE was calculated from ANPP and cumulative water supply. Bare soil evaporation (E) was calculated from climatic data and leaf area dynamics, and percolation (D) and transpiration (T) were estimated with HYDRUS-1D. Water-use efficiency (WUE, ANPP / (T + D)) and transpiration efficiency (TE, ANPP / T) were calculated. Resource availability had pronounced effects on the water-use efficiency of semiarid grassland. RUE, WUE, and TE all decreased under irrigated compared to rainfed conditions and were significantly increased with N fertilizer application at both levels of water supply. While the irrigation effect on parameters of water-use efficiency were accordingly reflected in stable carbon isotope signatures, N supply resulted in significantly less negative δ¹³C-values under rainfed but not irrigated conditions. It is concluded, that spatial or temporal gradients in resource availability have pronounced effects on the water-use efficiency of semiarid grassland. The decrease of water use-efficiency under high water supply was related to differences in TE and not to a relative increase of unproductive water loss. Carbon isotope discrimination was highly correlated with WUE and TE, but can be a poor predictor of RUE.     

呼伦贝尔草原生物量变化及其与环境因子的关系

根据呼伦贝尔草原大范围草地生物量的调查和实验数据,分析了该地区草地生物量的动态变化规律及其与环境因子的关系。结果表明,沿着环境梯度,不同区域草地生物量差异显著,其变化与水分、温度变化关系不显著,与0～20cm土层的土壤有机碳含量呈正相关,而与土壤容重呈负相关。逐步多元回归表明,土壤有机碳是制约生物量变化的主要因素,可能是当地草地利用方式使土壤养分成为制约草地植物生长的限制因子,从而影响草地生物量。     

Changes in some soil properties induced by re-conversion of cropland into grassland in the semiarid steppe zone of Inner Mongolia, China

Aims

“Grain for Green Program” (GGP), i.e., re-conversion of cropland into forest or grassland, initiated by Chinese government has a profound impact on mitigating environmental degradation. The objectives of this study were to assess the changes of some soil properties during the processes of re-conversion from cropland to grassland over time in the semiarid steppe region of north China.

Methods

Two sites with different ages of re-conversion were selected for measurements of organic matter (SOM), total nitrogen (TN) and phosphorus (TP), bulk density (BD) and grain size distribution. Saturated hydraulic conductivity was determined by the constant hydraulic head method and unsaturated hydraulic conductivity by disc infiltrometer at tensions of 30, 60 and 150 mm. Soil water content was measured using the gravimetric method. Wetting front depths in the soil after rainfall were also recorded at the study sites.

Results

Natural grasslands had higher belowground biomass than re-converted grasslands. Re-converted grasslands had lower SOM and TN at depths of 0–20 cm and higher saturated hydraulic conductivity at depths of 0–10 cm than natural grassland. The natural grassland soils had higher soil water contents in the surface soil (0–20 cm) and lower soil water contents at deeper depths than re-converted grassland soils. Soil aggregate stability reached the natural steppe level 12 years after re-conversion.

Conclusions

The recovery of soil properties after GGP appeared to be slow, and these properties did not return to natural grassland status before cultivation after 12 years of re-conversion.     

Tradeoffs between nitrogen- and water-use efficiency in dominant species of the semiarid steppe of Inner Mongolia

In water-limited environments, photosynthetic carbon gain and loss of water by transpiration are in a permanent tradeoff as both are contrarily regulated by stomata conductance. In semiarid steppe grasslands water limitation may covary with nitrogen limitation. Steppe grassland species are capable of optimizing their use of limiting resources, water and nitrogen, but regulation is still poorly understood. In a two-year experiment with addition of water (irrigation simulating a wet year) and nitrogen (0, 25, and 50 kg urea-N?ha^?1) we assessed intrinsic water use efficiency (WUE_i), nitrogen use efficiency (NUE), and related plant functional traits (PFTs) of four dominant C₃ species (Leymus chinensis, Agropyron cristatum, Stipa grandis, and Artemisia frigida). Water and N fertilizer supplementation significantly increased plant primary production, and N effect was more pronounced under irrigated conditions. Parallel with the responses of plant production, a strong tradeoff between WUE_i and NUE was detected: water supply increased NUE but decreased WUE_i, whereas N addition slightly increased WUE_i at the expense of NUE. This tradeoff occurred at the leaf level, and involved the responses of leaf N concentration and specific leaf area. WUE_i of species changed among treatments in a predictable manner by the parameter of leaf N content per area. Dominant plant species commonly achieved a higher utilization efficiency of the more limiting resource via altering PFTs, which was an important mechanism of adaptation to variable resource limitation in semiarid grasslands.     

Nutrient resorption responses to water and nitrogen amendment in semi-arid grassland of Inner Mongolia, China

Litterfall and fine root production is a major pathway for carbon and nutrient cycling in forest ecosystems. We investigated leaf litterfall, fine-root mass, production and turnover rate in the upper soil (0–30 cm) under four major tree species (Leucaena leucocephala, Acacia nilotica, Azadirachta indica, Prosopis juliflora) of the semi-arid region of India. All the four tree species showed an unimodal peak of leaf litterfall with distinct seasonality. Leucaena leucocephala and Acacia nilotica had maximum leaf litterfall between September and December while Azadirachta indica and Prosopis juliflora shed most of their leaves between February and May. Annual leaf litterfall of the four species ranged from 3.3 Mg ha^?1 (Leucaena leucocephala) to 8.1 Mg ha^?1 (Prosopis juliflora). Marked seasonal variations in amount of fine root biomass were observed in all the four tree species. Fine root production was maximum in Prosopis juliflora (171 g m^?2 y^?1) followed by Azadirachta indica (169 g m^?2 y^?1), Acacia nilotica (106 g m^?2 y^?1) and Leucaena leucocephala (79 g m^?2 y^?1). Fine root biomass showed a seasonal peak after the rainy season but fell to its lowest value during the winter and dry summer season. Fine root turnover rate ranged from 0.56 to 0.97 y^?1 and followed the order Azadirachta indica > Leucaena leucocephala > Prosopis juliflora > Acacia nilotica. The results of this study demonstrated that Prosopis juliflora and Azadirachta indica had greater capability for maintaining site productivity as evidenced from greater leaf litterfall and fine root production.     

Effects of root-feeding nematodes on aboveground net primary production in a North American grassland

The impact of nematode consumption on aboveground net primary production (ANPP) was addressed by treating two areas of mixed-grass prairie with the nematicide carbofuran. Monthly assessments of shoot biomass and nematode populations were made from each treated and control site for two growing seasons. Seasonal mean density of root-feeding nematodes was reduced approximately 82% by carbofuran. ANPP was significantly greater in the treated plots for both seasons in Site 1 but only for the second year in Site 2. Increases in ANPP averaged 51% in Site 1 and 26% in Site 2. It was estimated that root-feeding nematodes reduced ANPP by 16 times more than they consumed.     

Resource capture, biomass allocation and growth in herbaceous plants

Plant species differ widely in their rate of biomass production, even when grown under optimal conditions. A key question concerns the extent to which these growth rates correlate with the uptake of carbon and nitrogen and with the biomass allocation between leaves and roots. Recent data show that the answer to this question differs for mono- and dicotyledons, and that more than biomass allocation, it is the ratio between the activities of leaves and roots that correlates with the growth rate of a plant.     

Genotypic effects of fertilization on seedling sweetgum biomass allocation,N uptake,and N use efficiency

Screening and selecting tree genotypes that are responsive to N additions and that have high nutrient use efficiencies can provide better genetic material for short-rotation plantation establishment. A pot experiment was conducted to test the hypotheses that (1) sweetgum ( Liquidambar styraciflua L.) families have different patterns in biomass production and allocation, N uptake, and N use efficiency (NUE), because of their differences in growth strategies, and (2) sweetgum families that are more responsive to N additions will also have greater nutrient use efficiencies. Seedlings from two half-sib families (F10022 and F10023) that were known to have contrasting responses to fertility and other stress treatments were used for an experiment with two levels of N (0 vs. 100 kg N/ha equivalent) and two levels of P (0 vs. 50 kg P/ha equivalent) in a split-plot design. Sweetgum seedlings responded to N and P treatments rapidly, with increases in both size and biomass production, and those responses were greater with F10023 than with F10022. Growth response to N application was particularly strong. N and P application increased the proportional allocation of biomass to leaves. Under increased N supply, P application increased foliar N concentration and content, as well as total N uptake by the seedlings. However, NUE was decreased by N addition and was higher in F10023 than in F10022 when P was not limiting. A better understanding of genotype by fertility interactions is important in selecting genotypes for specific site conditions and for optimizing nutrient use in forestry production.     

Spatial and temporal variability in the net primary production of grassland in China and its relation to climate factors

As an important component of the terrestrial carbon (C) cycle, variability in net primary productivity (NPP) plays a crucial role in the C input and accumulation in grasslands system. In this study, the spatial and temporal variability of grassland NPP in China during 2001–2010 and its relation to climate factors were analyzed by using a modified model of Carnegie–Ames–Stanford Approach based on the Comprehensive and Sequential Classification System. The results show that monthly grassland NPP increases from January to July. While the seasonal variability of NPP indicates peak productivity in summer. Annual mean grassland NPP follows a significant increasing trend with fluctuation from 2001 to 2010. The spatial pattern of grassland NPP shows increasing gradients from the west to the east and from the north to the south of China. Annual NPP differs significantly among different grassland types, with the highest NPP in the grassland distributed in sub-tropical perhumid evergreen broad leaved forest and tropical-perhumid rain forest. Time-lag correlation analysis at the monthly scale shows that grassland NPP responded more rapidly to changes in temperature than to precipitation. Among the climate factors, grassland NPP shows the strongest correlation at 1-month lag with moisture index K. There is a significant positive correlation between seasonal NPP and K. The seasonal NPP is significantly correlated with >0 °C annual cumulative temperature. The highest and the lowest NPP sensitivity to precipitation, K, and temperature were observed in the grassland distributed in tropical forest and semi-desert. The results indicate a complex mechanism of climate factors that control grassland C sequestration in terrestrial ecosystems.     

Testing mechanisms of N-enrichment-induced species loss in a semiarid Inner Mongolia grassland: critical thresholds and implications for long-term ecosystem responses

The increase in nutrient availability as a consequence of elevated nitrogen (N) deposition is an important component of global environmental change. This is likely to substantially affect the functioning and provisioning of ecosystem services by drylands, where water and N are often limited. We tested mechanisms of chronic N-enrichment-induced plant species loss in a 10-year field experiment with six levels of N addition rate. Our findings on a semi-arid grassland in Inner Mongolia demonstrated that: (i) species richness (SR) declined by 16 per cent even at low levels of additional N (1.75 g N m^–2 yr⁻¹), and 50–70% species were excluded from plots which received high N input (10.5–28 g N m⁻² yr⁻¹); (ii) the responses of SR and above-ground biomass (AGB) to N were greater in wet years than dry years; (iii) N addition increased the inter-annual variations in AGB, reduced the drought resistance of production and hence diminished ecosystem stability; (iv) the critical threshold for chronic N-enrichment-induced reduction in SR differed between common and rare species, and increased over the time of the experiment owing to the loss of the more sensitive species. These results clearly indicate that both abundance and functional trait-based mechanisms operate simultaneously on N-induced species loss. The low initial abundance and low above-ground competitive ability may be attributable to the loss of rare species. However, shift from below-ground competition to above-ground competition and recruitment limitation are likely to be the key mechanisms for the loss of abundant species, with soil acidification being less important. Our results have important implications for understanding the impacts of N deposition and global climatic change (e.g. change in precipitation regimes) on biodiversity and ecosystem services of the Inner Mongolian grassland and beyond.     

Plant biomass and net production ofanogeissus latifolia Wall. in forests of semiarid zone of rajasthan (india)

A. latifolia grown in the Borimalan forest block in Prasad range (24°11′N and 73°42′ E) exerts clear positive correlations between CBH (circumference breast height)and number of growth rings of bole and branches, tree height, total biomass and leaf area. The net above-ground biomass is 3.95 × 10⁴ kg ha^-1. The average increment in non-photosynthetic (trunk + branch) biomass shows two peaks, the lower peak at 11–16 growth ring period, and the higher one at 34–36 growth ring period. The ratio of leaf dry weight/leaf area is16.3 to 34.8 mg cm^-2, the ratios between shoot net production: leaf weight and leaf area are1.5 g per g and 212 g m^-2 respectively.     

Influence of floristic composition on the net primary production and dry matter turnover in a tropical grassland

Plant biomass, net primary productivity and dry matter turnover were studied in a grassland situated in a tropical monsoonal climate at Kurukshetra, India (29°58′N, 76°51′E). Based on differences in vegetation in response to microrelief, three stands were distinguished on the study site. The stand I was dominated by Sesbania bispinosa, stand II represented mixed grasses and stand III was dominated by Desmostachya bipinnata. Floristic composition of the three stands revealed the greatest number of species on stand II (75). The study of life form classes indicated a thero-cryptophytic flora. The biomass of live shoots in all the three stands attained a maximum value in September (424–1921 g m^-2) and below ground plant biomass in November (749–1868 g m^-2). The annual above ground net primary production was greatest on stand I (2143 g m^-2) and lowest on stand II (617 g m^-2). The rate of production was highest during the rainy season (15.34 to 3.18 g m^-2 day^-2). Below ground net production ranged from 1592 to 785 g m^-2 y^-2 and the rates were high in winter and summer seasons. Total annual net primary production was estimated to be 3141, 1403, 2493 and 2134 g m^-2 on stands I, II, III and on the grassland as a whole, respectively. The turnover of total plant biomass plus below ground biomass indicated almost a complete replacement of phytomass within the year. The system transfer functions showed greater transfer of material from total net primary production to the shoot compartment during rainy season and to the root compartment during winter and summer seasons.     

The effects of spacing on growth, morphology and biomass production and allocation in two hybrid poplar clones growing in the boreal region of Canada

Intra-clonal competition was studied in young hybrid poplar plantations to assess the effects of spacing on growth, biomass production and allocation, and morphological characteristics of above- and below-ground tree parts. Three spacings were used as whole-plots (1 × 1 m, 3 × 3 m and 5 × 5 m), with two hybrid poplar clones as subplots (BT747, Populus balsamifera L. × P. trichocarpa Torr. & Gray; MB915, P. maximowiczii A. Henry × P. balsamifera L.) in a split-plot design. After six growing seasons, diameter at breast height (dbh) increased by about 120% from the 1 × 1 m to the 5 × 5 m spacing for clone MB915, while there was no significant change in dbh for the other clone. The effect of spacing on height growth was opposite for the clones; it increased by about 175% from the narrowest to the widest spacing for clone MB915, while it decreased by about 27% for clone BT747. Estimates of above-ground biomass production after six growing seasons were significantly reduced with increasing spacing, with 29.6, 4.9 and 3.2 MgDM ha⁻¹ on average from the narrowest to the widest spacing. Branch traits and the vertical distribution of leaf area were the most affected by spacing for both clones, while live crown ratio and percentage of syllepsis did not change. Spacing also affected proportions of biomass allocated to stem, leaves, and branches, but allocation to roots did not change.     

Biomass allocation, relative competitive ability and water use efficiency of two dominant species in semiarid Loess Plateau under water stress

A better understanding of the growth and interspecific competition of native dominant species under water stress should aid in prediction of succession in plant communities. In addition, such research would guide the selection of appropriate conservation and agricultural utilization of plants in semiarid environments that have not been very well characterized. Biomass production and allocation, relative competitive ability and water use efficiency of one C₄ herbaceous grass (Bothriochloa ischaemum) and one C₃ leguminous subshrub (Lespedeza davurica), both important species from the semiarid Loess Plateau of China, were investigated in a pot-cultivation experiment. The experiment was conducted using a replacement series design in which B. ischaemum and L. davurica were grown with twelve plants per pot, in seven combinations of the two species (12:0, 10:2, 8:4, 6:6, 4:8, 2:10, and 0:12). Three levels of water treatments included sufficient water supply (HW), moderate water stress (MW) and severe water stress (LW). These treatments were applied after seedling establishment and remained until the end of the experiment. Biomass production and its partitioning, and transpiration water use efficiency (TWUE) were determined at the end of the experiment. Interspecific competitive indices (competitive ratio (CR), aggressiveness (A) and relative yield total (RYT)) were calculated from the dry weight for shoots, roots and total biomass. Water stress decreased biomass production of both species in monoculture and mixture. The growth of L. davurica was restrained in their mixtures for each water treatment. L. davurica had significantly (P < 0.05) greater root:shoot allocation than B. ischaemum for each water treatment and proportion within the replacement series. Aggressiveness (A) values for B. ischaemum with respect to L. davurica were negative only at the proportions of B. ischaemum to L. davurica being 8:4 and 10:2 in LW treatment. B. ischaemum had a significantly (P < 0.05) higher CR value under each water treatment, and water stress considerably reduced its relative CR while increased that of L. davurica. RYT values of the two species indicated some degree of resource complimentarity under both water sufficient and deficit conditions. The results suggest that it is advantageous for growing the two species together to maximize biomass production, and the suggested ratio was 10:2 of B. ischaemum to L. davurica because of significantly higher (P < 0.05) RYT and TWUE under low water availability condition.     

Impacts of caterpillar disturbance on forest net primary production estimation in China

The study described the uncertainties of NPP estimation due to larch defoliation by caterpillars during 2001–2004 in Northeastern China, which is an order of magnitude larger in area and severe than all previous recorded outbreaks. Using “representative needles” collected in the field, we related spectral responses and photosynthetic contents of needles to ecophysiological parameters in NPP estimation for different degrees of caterpillar damage. It is concluded that locations of reflection peaks at or near green red near-infrared wavelength moving toward long-wave direction with aggravated damage degree, which is useful to earlier prediction of caterpillar damage. Contents of chlorophyll-a reduced by 40.26, 66.29 and 86.88% for light, moderate and severe damage samples respectively compared with healthy, chlorophyll-b reduced by 20.79, 52.67 and 82.39%, and carotenoid reduced by 9.80, 45.06 and 73.15%. Estimated NDVI values decreased by 9.89–31.08, 38.16–47.47 and 61.90–66.71%, respectively. It results to the decrease of LAI by 39.76–74.34, 85.85–93.64 and 94.61–98.51%, and reduction of FPAR and APAR by 10.16–37.47, 39.21–57.24 and 63.60–80.44%. In addition, the increased minimum visible reflectance during 580–680 nm leads to the decrease of estimated aboveground biomass by 14.62, 48.86 and 56.84%, and reduction of estimated autotrophic respiration by 10.32, 39.11 and 46.95%. Ultimately, the caterpillar outbreak results to the directly underestimation of NPP by 10.01–64.63, 39.32–75.37 and 80.26–100% for light, moderate and severe samples compared with healthy. Furthermore, such impacts of forest insect disturbances should be accounted for in NPP modeling to reduce the uncertainties of estimation.