Tree species richness as an important biotic factor regulates the soil phosphorus density in China's mature natural forests.

Tree species richness has been recognized as an underlying driving factor for regulating soil phosphorus (P) status in many site-specific studies. However, it remains poorly understood whether this is true at broad scales where soil P strongly rely on climate, soil type and vegetation type. Here, based on the data of 946 mature natural forest sites from a nationwide field survey in China, we analyzed the impact of tree species richness on soil P density of China's mature natural forests (deciduous coniferous forest, DCF; evergreen coniferous forest, ECF; deciduous broad-leaved forest, DBF; evergreen broad-leaved forest, EBF; and mixed coniferous and broad-leaved forest, MF). Our results showed that tree species richness had a negative effect on soil P density in China's mature natural forests. The Random Forest regression model showed that the relative importance of tree species richness to soil P density was second only to the climate factors (mean annual temperature, MAT; mean annual precipitation, MAP). In addition, the structural equation model (SEM) results showed that the goodness fit of SEM increased when the tree species richness was included into the model. These results suggested that tree species richness was an important factor in regulating the China's mature natural forests soil P density. Furthermore, the SEM results showed that the decreased soil P density was related to the increase in ANPP and the decrease in litter P concentration induced by tree species richness. This result indicates that tree species richness could facilitate plant P absorption and inhibit plant P return into the soil, and thus reducing the soil P density in China's mature natural forests. In conclusion, we found tree species richness was an important biotic factor in regulating soil P density at broad scales, which should be fully considered in Earth models that represent P cycle.

Warming drives sustained plant phosphorus demand in a humid tropical forest.

Phosphorus (P) is often one of the most limiting nutrients in highly weathered soils of humid tropical forests and may regulate the responses of carbon (C) feedback to climate warming. However, the response of P to warming at the ecosystem level in tropical forests is not well understood because previous studies have not comprehensively assessed changes in multiple P processes associated with warming. Here, we detected changes in the ecosystem P cycle in response to a 7-year continuous warming experiment by translocating model plant-soil ecosystems across a 600-m elevation gradient, equivalent to a temperature change of 2.1°C. We found that warming increased plant P content (55.4%) and decreased foliar N:P. Increased plant P content was supplied by multiple processes, including enhanced plant P resorption (9.7%), soil P mineralization (15.5% decrease in moderately available organic P), and dissolution (6.8% decrease in iron-bound inorganic P), without changing litter P mineralization and leachate P. These findings suggest that warming sustained plant P demand by increasing the biological and geochemical controls of the plant-soil P-cycle, which has important implications for C fixation in P-deficient and highly productive tropical forests in future warmer climates.

Natural forests promote phosphorus retention in soil.

Soil phosphorus (P) availability often limits plant productivity. Classical theories suggest that total P content declines at the temporal scale of pedogenesis, and ecosystems develop toward the efficient use of scarce P during succession. However, the trajectory of ecosystem P within shorter time scales of succession remains unclear. We analyzed changes to P pools at the early (I), middle (II), and late (III) stages of growth of plantation forests (PFs) and the successional stages of natural forests (NFs) at 1969 sites in China. We found significantly lower P contents at later growth stages compared to earlier ones in the PF (p < .05), but higher contents at late successional stages than in earlier stages in the NF (p < .05). Our results indicate that increasing P demand of natural vegetation during succession, may raise, retain, and accumulate P from deeper soil layers. In contrast, ecosystem P in PF was depleted by the more rapidly increasing demand outpacing the development of a P-efficient system. We advocate for more studies to illuminate the mechanisms for determining the divergent changes, which would improve forest management and avoid the vast degradation of PF ecosystems suffering from the ongoing depletion of P.

Responses of soil buffering capacity to acid treatment in three typical subtropical forests.

Elevated anthropogenic acid deposition can significantly affect forest ecosystem functioning by changing soil pH, nutrient balance, and chemical leaching and so on. These effects generally differ among different forests, and the dominant mechanisms for those observed responses often vary, depending on climate, soil conditions and vegetation types. Using soil monoliths (0-40cm) from pine forest (pioneer), coniferous and broadleaved mixed forest (transitional) and broadleaved forest (mature) in southern China, we conducted a leaching experiment with acid treatments at different pH levels (control: pH≈4.5; pH=3.5; pH=2.5). We found that pH3.5 treatment significantly reduced dissolved organic carbon (DOC) concentrations in leachate from the pioneer forest soil. pH2.5 treatment significantly increased concentrations of NO3(-), SO4(2-), Ca(2+), Mg(2+), Al(3+), Fe(3+) and DOC in leachate from the pioneer forest soil, and also concentrations of NO3(-), SO4(2-), Mg(2+), Al(3+), Fe(3+) and DOC in leachate from the transitional forest soil. All acid treatments had no significant effects on concentrations of these chemicals in leachate from the mature forest soil. The responses can be explained by the changes in soil pH, acid neutralizing capacity (ANC) and concentrations of Al and Fe. Our results showed that acid buffering capacity of the pioneer or transitional forest soil was lower than that of the mature forest soil. Therefore preserving mature forests in southern China is important for reducing the adverse impacts of high acid deposition on stream water quality at present and into the future.

The mechanism for exclusion of Pinus massoniana during the succession in subtropical forest ecosystems: light competition or stoichiometric homoeostasis?

Competition for light has traditionally been considered as the main mechanism for exclusion of Pinus massoniana during succession in subtropical forest ecosystems. However, both long-term inventories and a seedling cultivation experiment showed that growth of mature individuals and young seedlings of P. massoniana was not limited by available light, but was strongly influenced by stoichiometric homoeostasis. This is supported by the results of homoeostatic regulation coefficients for nitrogen (HN) and phosphorus (HP) estimated using the measured data from six transitional forests across subtropical China. Among three dominant tree species in subtropical forests, P. massoniana and Castanopsis chinensis had the lowest values of HP and HN, respectively. Therefore P. massoniana cannot survive in the advanced stage due to soil phosphorus limitation and C. chinensis cannot successfully grow in the pioneer stage due to soil nitrogen limitation. Our results support that stoichiometric homeostasis is the main reason for gradual exclusion of P. massoniana from the transitional forest and the eventual elimination from the advanced forest during the subtropical forest succession. Therefore greater attention should be paid to stoichiometric homeostasis as one of the key mechanisms for species exclusion during forest succession.

Available phosphorus in forest soil increases with soil nitrogen but not total phosphorus: evidence from subtropical forests and a pot experiment.

This paper aims to establish evidence for available phosphorous (AP) binding with total nitrogen (N) in subtropical forest soils. Soil organic carbon (SOC), total N, total phosphorous (P) and AP concentration were measured for three contrasting forest types in southern China: Masson pine forest (MPF), coniferous and broadleaved mixed forest (CBMF) and monsoon evergreen broadleaved forest (MEBF). A pot experiment with N addition was conducted to confirm the dominant factor to affect on soil AP concentration. The results showed that mean soil total N concentration in 0-10 cm soil layer was 440 ± 50 for MPF, 840 ± 80 for CBMF and 1020 ± 50 mg kg(-1) for MEBF, respectively. The mean soil AP concentration in 0-10 cm soil layer was 2.67 ± 0.87 for MPF, 2.65 ± 0.58 for CBMF, 4.10 ± 0.29 mg kg(-1) for MEBF, respectively. The soil total N concentration could explain about 70% of the variations in soil AP concentration in the top 20 cm soil layers in the three forest types. A pot experiment with N addition also showed an increase of AP concentration from 2.56 to 5.63 mg kg(-1), when N addition increased from 5 g to 17 g NH4NO3. Our results therefore suggested that N addition significantly increased soil AP concentration, which might be beneficial for stabilizing the net primary production of subtropical forests that were limited by soil AP. This finding may provide a theory basis for tropical and subtropical forests management.

Synthesis of ethyleneoxide modified 3-carboranyl thymidine analogues and evaluation of their biochemical, physicochemical, and structural properties.

Eleven 3-carboranyl thymidine analogues (3CTAs) containing highly hydrophilic and flexible ethyleneoxide moieties were synthesized as potential agents for boron neutron capture therapy (BNCT) and their biochemical and physicochemical properties were evaluated. Based on specific structural features, this library of 3CTAs was divided into three subgroups. The first group contained 3CTAs with 1-4 ethyleneoxide units between the thymidine (Thd) scaffold and a carborane cluster. The second group of 3CTAs contained a pentylene spacer between Thd and the carborane and 2-4 ethyleneoxide units additionally attached to the carborane cluster. The third group contained three 3CTAs all with pentylene spacers and four ethylene units but with different carborane cages. The ethyleneoxide modified 3CTAs were good substrates of thymidine kinase 1 (TK1) and poor substrates of human mitochondrial thymidine kinase 2 (TK2) as determined in phosphoryl transfer assays. In the first group of 3CTAs, all the compounds were efficiently phosphorylated regardless of varying spacer lengths (37-42% of the activity of Thd). The second group of 3CTAs was less effectively phosphorylated (17-26% of the activity of Thd) probably due to a less favorable sterical orientation of Thd within the active site of TK1 and/or an increased lipophilicity compared with the first group. In the third group of structural isomers, no significant differences in phosphorylation rates were observed (17-25%). A structure-function hypothesis explaining these results is presented.