We tested for the effects of PM2.5 pollution and climate change on satellite-observed spring greenness (NDVI, LAI, and SIF) through rigorous analysis of ground and satellite data (Supplementary Materials and Methods). Before that, we first examined and minimized the attenuation effects of PM2.5 pollution on the greenness signals of satellite observation (Supplementary Figs. 2 and 3). For the global scale, spatially explicit results for NDVI, LAI and SIF using satellite-based PM2.5 pollution were provided (Fig. 1 B-D). PM2.5 pollution led to reduced spring greening in 59.7-64.9% of the studied area, with 12.2-14.3% being significant. In comparison, PM2.5 pollution led to increased spring greening in ~38% of regions (e.g., west Australia, North Africa, high lands in the northern Europe), with only ~4% being statistically significant. Consistent results were observed for the USA, Europe and China where ground monitoring networks have been well established. The ground-sourced observations demonstrate that PM2.5 pollution were associated with decreased spring greening, with median standard sensitives for NDVI, LAI and SIF of -0.33, -0.36 and -0.38, respectively (Fig. 1 A).
We further conducted analyses to separate the greening effects of CO2, temperature, precipitation, vapor pressure deficit (VPD), and PM2.5 (Fig. 1 E-F). Stepwise regression analysis confirmed the dominant negative impacts of PM2.5 pollution on spring greening at site and global scales, the extent of which was comparable to the inhibiting effects of vapor pressure deficit (Extended Data Figs. 2 and 3). Given the potential multicollinearity of driving factors, we also used partial correlation and ridge regression analyses and found consistently and dominantly negative effects of PM2.5 pollution on spring greening (Extended Data Fig. 4 and Supplementary Fig. 4). Overall, these findings reveal that PM2.5 pollution inhibits photosynthesis and offsets the ongoing trends in land surface greening globally.
To study the underlying mechanisms that drive the observed reductions in spring vegetation activity in response to PM2.5 pollution, we conducted experiments on the leaf morphology of 15 widely-distributed tree species. We found that PM2.5 can adhere to the leaf surface to varying degrees (Supplementary Fig. 5), potentially causing blockages and damage to leaf stomata, as observed through scanning electron microscopy (Fig. 2A1-A15). To further investigate the effects of PM2.5 on gas exchange and photosynthesis, including stomatal size, density and conductance, transpiration rate, chlorophyll content, maximum CO2 assimilation rate, potential photosynthetic capacity (Fv/Fm), and photosynthetic rate, we conducted a meta-analysis based on 233 records of both experimental and observational results across 104 plant species worldwide (Fig. 2B). Overall, PM2.5 exposure caused substantial reductions in stomatal size (−0.15, P < 0.01), stomatal conductance (−0.18, P < 0.01), chlorophyll content (−0.17, P < 0.01), transpiration rate (−0.27, P < 0.01), and photosynthetic rate (−0.18, P < 0.01). Similar results were found when separately analyzing the experimental and observational data (Supplementary Fig. 6). Notably, the significant decline in stomatal conductance and photosynthetic rate was identified as the key link between plant growth and PM2.5 pollution. Further analysis of the effects of PM2.5 on stomatal conductance using global flux measurements showed consistent results with the meta-analysis, indicating that increased PM2.5 caused significant (P < 0.05) decreases in stomatal conductance and vegetation productivity accordingly (Fig. 2C and Supplementary Figs. 7 and 8). In line with flux measurements, PM2.5 pollution could lower canopy stomatal conductance and SIF based on gridded data, supporting that PM2.5 effects on greening trends are linked with the gas exchange between air and the interior of leaf (Supplementary Fig. 9).
To gain deeper insights into the underlying mechanisms that drive the correlation between PM2.5 pollution and spring greening, we explored potential biogeophysical and biogeochemical paths for the correlation (Fig. 3). We found that elevated levels of PM2.5 greatly reduced the amount of photosynthetically active radiation (PAR), with 64.7% of the grids showing a negative PM2.5-PAR correlation (16.2% of which were significant). In line with gridded data analysis, flux measurements confirmed the negative impacts of PM2.5 on PAR (Supplementary Fig. 10). This adverse effect on photosynthesis led to substantial declines in the maximum rate of carboxylation (VCmax), a key indicator of leaf photosynthetic capacity. This was evidenced by 64.1% negative correlations (12.3% significant) compared to only 35.9% positive correlations (3.6% significant). Since VCmax generally showed a positive correlation with SIF (62.0% positive vs. 1.2% negative, P < 0.05), higher PM2.5 levels counteracted the process of spring greening (Fig. 3A). Similar trends were observed for LAI. Nonetheless, certain regions exhibited increased spring greening with higher PM2.5. In these areas, PM2.5 raised the fraction of diffuse radiation (PARdiff/PAR) and nitrogen deposition (Ndeposition). Our structural equation models supported the hypothesis that PM2.5 decreased radiation, thereby reducing VCmax and LAI, but increased the fraction of diffuse radiation and Ndeposition (Fig. 3 C-H). Increased PM2.5 concentration could also lower the ambient ozone (O3) levels in spring due to lower atmospheric radiation, potentially undermining vegetation photosynthetic activities (Extended data Fig. 5). We also tested the impact of PM2.5 on air temperature and found no dominant relationship (Supplementary Fig. 11), suggesting PM2.5 pollution effect was not determined by air temperature regulation.
In a last step, we used the output from the TRENDY project to test the potential of 16 state-of-art terrestrial ecosystem models to reproduce the effects of PM2.5 on gross primary productivity (GPP), a productivity-based indicator of greening (Supplementary Table 5). Overall, the ecosystem models captured the widespread and negative effects of PM2.5 (Fig. 4A and Supplementary Fig. 12). The standard sensitivity of GPP to PM2.5 across the models was -0.15 ± 0.05, which is comparable with the inhibiting effects of VPD (Fig. 4B). However, the large biases of PM2.5 effects, indicated by relatively high standard deviation of PM2.5 sensitivity among models, were detected in the central of Europe and the eastern of America (Fig. 4C), suggesting the large inconsistency and limitation of model projections. Pixel-to-pixel comparison of PM2.5 sensitivities between satellite observations and model projections suggests an incorrect representation of PM2.5 effects on vegetation greening in nearly 34% of areas (Fig. 4D), highlighting the need for incorporating PM2.5 effects into future model improvement.