Global change progressively increases foliar nitrogen-phosphorus ratios in China's subtropical forests.

Globally increased nitrogen (N) to phosphorus (P) ratios (N/P) affect the structure and functioning of terrestrial ecosystems, but few studies have addressed the variation of foliar N/P over time in subtropical forests. Foliar N/P indicates N versus P limitation in terrestrial ecosystems. Quantifying long-term dynamics of foliar N/P and their potential drivers is crucial for predicting nutrient status and functioning in forest ecosystems under global change. We detected temporal trends of foliar N/P, quantitatively estimated their potential drivers and their interaction between plant types (evergreen vs. deciduous and trees vs. shrubs), using 1811 herbarium specimens of 12 widely distributed species collected during 1920-2010 across China's subtropical forests. We found significant decreases in foliar P concentrations (23.1%) and increases in foliar N/P (21.2%). Foliar N/P increased more in evergreen species (22.9%) than in deciduous species (16.9%). Changes in atmospheric CO2 concentrations ( P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ ), atmospheric N deposition and mean annual temperature (MAT) dominantly contributed to the increased foliar N/P of evergreen species, while P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ , MAT, and vapor pressure deficit, to that of deciduous species. Under future Shared Socioeconomic Pathway (SSP) scenarios, increasing MAT and P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ would continuously increase more foliar N/P in deciduous species than in evergreen species, with more 12.9%, 17.7%, and 19.4% versus 6.1%, 7.9%, and 8.9% of magnitudes under the scenarios of SSP1-2.6, SSP3-7.0, and SSP5-8.5, respectively. The results suggest that global change has intensified and will progressively aggravate N-P imbalance, further altering community composition and ecosystem functioning of subtropical forests.

Nitrogen and phosphorus addition promote invasion success of invasive species via increased growth and nutrient accumulation under elevated CO2.

In the context of the resource allocation hypothesis regarding the trade-off between growth and defence, compared with native species, invasive species generally allocate more energy to growth and less energy to defence. However, it remains unclear how global change and nutrient enrichment will influence the competition between invasive species and co-occurring native species. Here, we tested whether nitrogen (N) and phosphorus (P) addition under elevated CO2 causes invasive species (Mikania micrantha and Chromolaena odorata) to produce greater biomass, higher growth-related compounds and lower defence-related compounds than native plants (Paederia scandens and Eupatorium chinense). We grew these native and invasive species with similar morphology with the addition of N and P under elevated CO2 in open-top chambers. The addition of N alone increased the relative growth rate (RGR) by 5.4% in invasive species, and its combination with P addition or elevated CO2 significantly increased the RGR of invasive species by 7.5 or 8.1%, respectively, and to a level higher than that of native species (by 14.4%, P < 0.01). Combined N + P addition under elevated CO2 decreased the amount of defence-related compounds in the leaf, including lipids (by 17.7%) and total structural carbohydrates (by 29.0%), whereas it increased the growth-related compounds in the leaf, including proteins (by 75.7%), minerals (by 9.6%) and total non-structural carbohydrates (by 8.5%). The increased concentrations of growth-related compounds were possibly associated with the increase in ribulose 1,5-bisphosphate carboxylase oxygenase content and mineral nutrition (magnesium, iron and calcium), all of which were higher in the invasive species than in the native species. These results suggest that rising atmospheric CO2 concentration and N deposition combined with nutrient enrichment will increase the growth of invasive species more than that of native species. Our result also suggests that invasive species respond more readily to produce growth-related compounds under an increased soil nutrient availability and elevated CO2.

Effects of elevated CO2 concentration and nitrogen addition on foliar phosphorus fractions of Mikania micranatha and Chromolaena odorata under low phosphorus availability.

Invasive plants rapidly spread in habitats with low soil phosphorus (P) availability and have triggered a sharp decline in the diversity of native species. However, no studies have explored how widespread invasive species acclimate to low soil P availability via changing foliar P fractions, especially under elevated atmospheric CO2 concentrations ([CO2 ]) and nitrogen (N) deposition. Here, an open-top chamber experiment was conducted to explore the effect of nutrient addition and elevated [CO2 ] on leaf traits and foliar functional P fractions (i.e., Pi, metabolite P, lipid P, nucleic acid P, and residual P) of two aggressive invasive species (Mikania micranatha and Chromolaena odorata). We found that foliar N/P ratios were more than 20, and P addition significantly increased plant biomass. Both results indicated P-limited plant growth at our studied site. Elevated [CO2 ], N and N + P addition greatly increased plant biomass, photosynthetic rates, and photosynthetic P-use efficiency (PPUE) in invasive species, but PPUE decreased with increasing P addition. Nitrogen addition slightly decreased the concentration of leaf total P, decreased foliar residual P, but increased metabolite P concentrations in invasive species. Similar changes in foliar P fractions were found under N + P addition. Phosphorus addition increased foliar P concentrations, which was strongly correlated with an increase in metabolite P concentrations in invasive species. Elevated [CO2 ] alleviated these effects and increased PPUE. The present results suggest that future elevated [CO2 ] and N deposition allow the invasive species to acclimate to low soil P availability and support their successful invasion by greatly reducing P allocation to non-metabolite foliar P fractions (i.e., nucleic acid P and residual P) to meet their demand of metabolite P for photosynthesis and exhibit a high PPUE.

Latitudinal patterns of terrestrial phosphorus limitation over the globe.

Phosphorus limitation on terrestrial plant growth is being incorporated into Earth system models. The global pattern of terrestrial phosphorus limitation, however, remains unstudied. Here, we examined the global-scale latitudinal pattern of terrestrial phosphorus limitation by analysing a total of 1068 observations of aboveground plant production response to phosphorus additions at 351 forest, grassland or tundra sites that are distributed globally. The observed phosphorus-addition effect varied greatly (either positive or negative), depending significantly upon fertilisation regime and production measure, but did not change significantly with latitude. In contrast, phosphorus-addition effect standardised by fertilisation regime and production measure was consistently positive and decreased significantly with latitude. Latitudinal gradient in the standardised phosphorus-addition effect was explained by several mechanisms involving substrate age, climate, vegetation type, edaphic properties and biochemical machinery. This study suggests that latitudinal pattern of terrestrial phosphorus limitation is jointly shaped by macro-scale driving forces and the fundamental structure of life.

Global meta-analysis shows pervasive phosphorus limitation of aboveground plant production in natural terrestrial ecosystems.

Phosphorus (P) limitation of aboveground plant production is usually assumed to occur in tropical regions but rarely elsewhere. Here we report that such P limitation is more widespread and much stronger than previously estimated. In our global meta-analysis, almost half (46.2%) of 652 P-addition field experiments reveal a significant P limitation on aboveground plant production. Globally, P additions increase aboveground plant production by 34.9% in natural terrestrial ecosystems, which is 7.0-15.9% higher than previously suggested. In croplands, by contrast, P additions increase aboveground plant production by only 13.9%, probably because of historical fertilizations. The magnitude of P limitation also differs among climate zones and regions, and is driven by climate, ecosystem properties, and fertilization regimes. In addition to confirming that P limitation is widespread in tropical regions, our study demonstrates that P limitation often occurs in other regions. This suggests that previous studies have underestimated the importance of altered P supply on aboveground plant production in natural terrestrial ecosystems.