Z. Y. Yuan

Fine Root Biomass, Production, Turnover Rates, and Nutrient Contents in Boreal Forest Ecosystems in Relation to Species, Climate, Fertility, and Stand Age: Literature Review and Meta-Analyses

Fine roots <2 mm in diameter play a key role in regulating the biogeochemical cycles of ecosystems and are important to our understanding of ecosystem responses to global climate changes. Given the sensitivity of fine roots, especially in boreal region, to climate changes, it is important to assess whether and to what extent fine roots in this region change with climates. Here, in this synthesis, a data set of 218 root studies were complied to examine fine root patterns in the boreal forest in relation to site and climatic factors. The mean fine root biomass in the boreal forest was 5.28 Mg ha−1, and the production of fine roots was 2.82 Mg ha−1 yr−1, accounting for 32% of annual net primary production of the boreal forest. Fine roots in the boreal forest on average turned over 1.07 times per year. Fine roots contained 50.9 kg ha−1 of nitrogen (N) and 3.63 kg ha−1 of phosphorous (P). In total, fine roots in the boreal forest ecosystems contain 6.1 × 107 Mg N and 4.4×106Mg P pools, respectively, about 10% of the global nutrients of fine roots. Fine root biomass, production, and turnover rate generally increased with increasing mean annual temperature and precipitation. Fine root biomass in the boreal forest decreased significantly with soil N and P availability. With increasing stand age, fine root biomass increased until about 100 years old for forest stands and then leveled off or decreased thereafter. These results of meta analysis suggest that environmental factors strongly influence fine root biomass, production, and turnover in boreal forest, and future studies should place a particular emphasis on the root-environment relationships.

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.

Tree species diversity increases fine root productivity through increased soil volume filling

Summary 1. Although fine roots (< 2 mm in diameter) account for a major share of the production of terrestrial ecosystems, diversity effects on fine root productivity and their mechanisms remain unclear. 2. We hypothesized that: (i) fine root productivity increases with tree species diversity, (ii) higher fine root productivity is a result of greater soil volume filling due to species-specific patterns of root placement and proliferation, and (iii) differences in fine root productivity and soil volume filling associated with tree species diversity are more pronounced in summer when plants are physiologically active and demand for water and nutrients is at its greatest. 3. We investigated the effects of tree species diversity on fine root productivity and soil volume filling of boreal forest stands that have grown naturally for 85 years on similar sites. 4. Annual fine root production was 19–83% higher in evenly mixed- than single-species-dominated stands, and increased with tree species evenness, but not tree species richness. Fine root biomass was higher in evenly mixed- than single-species-dominated stands in summer months, but not in spring or fall. Higher fine root productivity in evenly mixed- than single-species-dominated stands was realized by filling more soil volume horizontally and vertically in the forest floor in the mixtures of deep- and shallow-rooted species vs. the deeper mineral soil in the mixtures of deeprooted species. 5. Synthesis. Our results provide some of the first direct evidence for below-ground species complementarity in heterogeneous natural forests, by demonstrating that tree species evenness increases fine root productivity by filling/exploiting the soil environment more completely in space and time, driven by differences in the inherent rooting traits of the component species and variations of root growth within species.

Global-scale latitudinal patterns of plant fine-root nitrogen and phosphorus

Most water and essential soil nutrient uptake is carried out by fine roots in plants. It is therefore important to understand the global geographic patterns of fine-root nitrogen and phosphorus cycling. Here, by compiling plant root data from 211 studies in 51 countries, we show that live fine roots have low nitrogen (N) and phosphorus (P), but similar N:P ratios when compared with green leaves. The fine-root N:P ratio differs between biomes and declines exponentially with latitude in roots of all diameter classes. This is in contrast to previous reports of a linear latitudinal decline in green leaf N:P, but consistent with nonlinear declines in leaf litter N:P. Whereas the latitudinal N:P decline in both roots and leaves reflects collective influences of climate, soil age and weathering, differences in the shape of the response function may be a result of their different N and P use strategies.

Negative effects of fertilization on plant nutrient resorption

Plants in infertile habitats are thought to have a high rate of nutrient resorption to enable them reuse nutrients more efficiently than those in fertile habitats. However, there is still much debate on how plant nutrient resorption responds to nutrient availability. Here we used a meta-analysis from a global data set of 9703 observations at 306 sites from 508 published articles to examine the effects of nitrogen (N) and phosphorus (P) fertilization on plant foliar N and P concentrations and resorption efficiency. We found that N fertilization enhanced N concentration in green leaves by 27% and P fertilization enhanced green-leaf P by 73% on average. The N and P concentrations in senesced leaves also increased with respective nutrient fertilization. Resorption efficiencies (percentage of nutrient recovered from senescing leaves) of both N and P declined in response to respective nutrient fertilization. Combined N and P fertilization also had negative effects on both N and P resorption efficiencies. Whether nutrient resorption efficiency differs among plant growth types and among ecosystems, however, remains uncertain due to the limited sample sizes when analyzed by plant growth types or ecosystem types. Our analysis indicates that fertilization decreases plant nutrient resorption and the view that nutrient resorption is a critical nutrient conservation strategy for plants in nutrient-poor environments cannot be abandoned. The response values to fertilization presented in our analysis can help improve biogeochemical models.

A global analysis of fine root production as affected by soil nitrogen and phosphorus

Fine root production is the largest component of belowground production and plays substantial roles in the biogeochemical cycles of terrestrial ecosystems. The increasing availability of nitrogen (N) and phosphorus (P) due to human activities is expected to increase aboveground net primary production (ANNP), but the response of fine root production to N and P remains unclear. If roots respond to nutrients as ANNP, fine root production is anticipated to increase with increasing soil N and P. Here, by synthesizing data along the nutrient gradient from 410 natural habitats and from 469 N and/or P addition experiments, we showed that fine root production increased in terrestrial ecosystems with an average increase along the natural N gradient of up to 0.5 per cent with increasing soil N. Fine root production also increased with soil P in natural conditions, particularly at P < 300 mg kg−1. With N, P and combined N + P addition, fine root production increased by a global average of 27, 21 and 40 per cent, respectively. However, its responses differed among ecosystems and soil types. The global average increases in fine root production are lower than those of ANNP, indicating that above- and belowground counterparts are coupled, but production allocation shifts more to aboveground with higher soil nutrients. Our results suggest that the increasing fertilizer use and combined N deposition at present and in the future will stimulate fine root production, together with ANPP, probably providing a significant influence on atmospheric CO2 emissions.

Changes in nitrogen resorption traits of six temperate grassland species along a multi-level N addition gradient

Nitrogen (N) resorption from senescing leaves is an important mechanism of N conservation for terrestrial plant species, but changes in N-resorption traits over wide-range and multi-level N addition gradients have not been well characterized. Here, a 3-year N addition experiment was conducted to determine the effects of N addition on N resorption of six temperate grassland species belonging to three different life-forms: Stipa krylovii Roshev. (grass), Cleistogenes squarrosa (T.) Keng (grass), Artemisia frigida Willd. (semishrub), Melissitus ruthenica C.W.Wang (semishrub and N-fixer), Potentilla acaulis L. (forb) and Allium bidentatum Fisch.ex Prokh. (forb). Generally, N concentrations in green leaves increased asymptotically for all species. N concentrations in senescent leaves for most species (5/6) also increased asymptotically, except that the N concentration in senescent leaves of A. bidentatum was independent of N addition. N-resorption efficiency decreased with increasing N addition level only for S. krylovii and A. frigida, while no clear responses were found for other species. These results suggest that long-term N fertilization increased N uptake and decreased N-resorption proficiency, but the effects on N-resorption efficiency were species-specific for different temperate grassland species in northern China. These inter-specific differences in N resorption may influence the positive feedback between species dominance and N availability and thus soil N cycling in the grassland ecosystem in this region.

Increasing aridity, temperature and soil pH induce soil C-N-P imbalance in grasslands.

Due to the different degrees of controls exerted by biological and geochemical processes, climate changes are suggested to uncouple biogeochemical C, N and P cycles, influencing biomass accumulation, decomposition and storage in terrestrial ecosystems. However, the possible extent of such disruption in grassland ecosystems remains unclear, especially in China’s steppes which have undergone rapid climate changes with increasing drought and warming predicted moving forward in these dryland ecosystems. Here, we assess how soil C-N-P stoichiometry is affected by climatic change along a 3500-km temperate climate transect in Inner Mongolia, China. Our results reveal that the soil from more arid and warmer sites are associated with lower soil organic C, total N and P. The ratios of both soil C:P and N:P decrease, but soil C:N increases with increasing aridity and temperature, indicating the predicted decreases in precipitation and warming for most of the temperate grassland region could lead to a soil C-N-P decoupling that may reduce plant growth and production in arid ecosystems. Soil pH, mainly reflecting long-term climate change in our sites, also contributes to the changing soil C-N-P stoichiometry, indicating the collective influences of climate and soil type on the shape of soil C-N-P balance.

Indirect Methods Produce Higher Estimates of Fine Root Production and Turnover Rates than Direct Methods

The production and turnover of fine roots play substantial roles in the biogeochemical cycles of terrestrial ecosystems. However, the disparity among the estimates of both production and turnover, particularly due to technical limitations, has been debated for several decades. Here, we conducted a meta-analysis to compare published estimates of fine root production and turnover rates derived from different methods at the same sites and at the same sampling time. On average, the estimates of fine root production and turnover rates were 87% and 124% higher, respectively, by indirect methods than by direct methods. The substantially higher fine root production and turnover estimated by indirect methods, on which most global carbon models are based, indicate the necessity of re-assessing the global carbon model predictions for atmospheric carbon sequestration in soils as a result of the production and turnover of fine roots.

Soil water status influences plant nitrogen use: a case study

We studied differences in nitrogen use efficiency (NUE) among six species [Calamagrostis epigejos (L.) Roth., Carex duriuscula C.A. Mey., Phragmites communis (L.) Trin., Salix gordejevii Y.L. Chang, Salix cheilophila Schneid., and Typha minima Funk.] growing in two contrasting habitat types, i.e., a riverine wetland with high water supply and a riparian zone with low water supply. The two sites were different in soil water supply, but not in nitrogen supply. Here, NUE was defined as the total net primary production per unit nitrogen absorbed. There was no significant difference in NUE between the species growing in the riverine wetland (Carex duriuscula, P. communis, S. cheilophila, T. minima) and the species growing in the river bank (Carex duriuscula, Calamagrostis epigejos, P. communis, S. gordejevii). We further analyzed NUE as the product of the nitrogen productivity (A, the rate of dry matter production per unit of nitrogen in the plant) and the mean residence time of nitrogen (MRT, the period of time a unit of nitrogen is present in the plant). The species growing in the riverine wetland had larger A but lower MRT than the species growing in the river bank. There was an inverse relationship between A and MRT. Consequently, NUE was similar among species and habitats. These results suggested that environmental factors, such as soil water supply, can influence N use by plants.