Fertile island effects have rarely been compared between soil depths (Cao et al., 2021), particularly across large environmental gradients, despite the fact that effects may vary between different depths. In our study we showed strong fertile island effects at 54 dryland sites across the Tibetan Plateau but no differences between the two soil depths examined. We also demonstrated that the extent of the fertile island effect varied greatly among different soil fertility variables, with soil organic and inorganic carbon showing the highest and lowest effects at both depths, respectively. Overall, it is clear that our dryland shrub species have the capacity to accumulate carbon and nutrients at both the soil surface and at depth. The primary contributor of soil organic carbon to the soil is derived from plant litter (Dove et al., 2019). Plant roots additionally provide readily available exudates to nourish soil microbes, promoting their proliferation and conversion into organic carbon within the soil (Huang et al., 2000; Sokol & Bradford, 2019; Villarino et al., 2021). Therefore, it is not unexpected that we found a greater quantities of soil organic carbon beneath the plant canopies than the open areas. Carbonates formed during weathering are a major source of inorganic carbon in dryland ecosystems (Pan & Guo, 1999). Hence, changes in soil inorganic carbon were less affected by plants and that the fertile island effect was the lowest (Fig. 1).
In the surface layer, we observed a notable increase in the fertile island effects for soil organic carbon as precipitation seasonality increased. The surface soil layer experiences more rapid fluctuations in water conditions, i.e., it is frequently wet and dries out faster than to the deeper soil layers (Ebrahimi & Or, 2015). These varying wet-dry cycles in the upper layer may lead to an elevated soil fungal-to-bacterial ratio, thereby enhancing the capacity of the soil to retain nutrients (Gordon et al., 2008; Tecon & Or, 2016). Our structural equation modelling further revealed that organic carbon affected by the fertile island effect was directly impacted by precipitation seasonality. This link may be attributed to the structural and functional alterations that occur in microbial communities in response to wet-dry alternations (Gordon et al., 2008; Tecon & Or, 2016). We noted that the fertile island effects of total carbon increased with rising precipitation in the wettest quarter and increasing precipitation seasonality, but decreased with higher temperatures during the driest quarter. Greater precipitation seasonality appeared to intensify the magnitude of the fertile island effects for soil organic carbon, subsequently influencing the fertile island effects of total carbon. Moreover, the fertile island effect for inorganic carbon increased with increasing precipitation during the wettest quarter, but decreased with increasing temperature of the driest quarter. Higher precipitation levels in dryland soils tend to increase soil water content and microbial biomass (Manzoni et al., 2014; Nielsen & Ball, 2015; Sierra et al., 2017). Microorganisms, under these conditions, effectively decompose organic acids and matter into carbonate minerals, which ultimately contribute to inorganic carbon in the soil (Ferdush & Paul, 2021). The study area experiences an average temperature below freezing during the driest quarter, and snow cover provides thermal insulation that retards microbial metabolic activity. However, increasing temperatures, frequent freeze-thaw cycles and greater temperature fluctuations can have detrimental effects on surface soil microorganisms, ultimately inhibiting the organic carbon mineralization process into inorganic carbon (Campbell et al., 2005).
In the underlying layer of soil, it was observed that the fertile island effect for nitrogen increased with higher soil sand content, while the effect for phosphorus trended upwards with increasing vegetation greenness. In these deeper soil layers, the availability of oxygen is comparatively lower than in the surface layer, and the carbon substrate remains relatively stable, which limits the decomposition of carbon (Wang et al., 2022). Additionally, temperatures in deeper soil layers are less influenced by atmospheric temperature fluctuations compared to surface depths, which makes the fertile island effect of soil carbon less sensitive to climatic factors at this depth. In open areas, as soil sand content increases, the adsorption of nitrogen compounds by clay is inhibited, hindering the nitrogen mineralization process and resulting in reduced nitrogen content (Li et al., 2022). Furthermore, soils containing a greater proportion of sand are prone to experiencing nitrogen loss due to elevated levels of leaching and runoff (Dai et al., 2022; Zhao et al., 2022). Plants are the primary conduits for nitrogen entry into the soil (Chen et al., 2021). In areas with vegetation cover, plants act to reduce soil nitrogen loss through leaching and runoff (von Felten et al., 2009). Additionally, plant leaf and root litter make up a substantial source of soil organic nitrogen (Chen et al., 2022). Abundant nutrients derived from plants and the creation of diverse microbial habitats within plant communities can contribute to the promotion of soil microbial biomass and activity, leading to an acceleration in both nitrogen mineralization and immobilization rates (Hooper & Vitousek, 1998; Zak et al., 2003). These factors contribute to the observed increase in soil nitrogen content differences between subcanopy areas and bare sections. Consequently, the fertile island effect for total nitrogen and organic nitrogen exhibited an increase with coarser soil texture.
In the deeper soil layer, the fertile island effect for total phosphorus significantly increased with a rise in the value of the NDVI. A higher NDVI typically indicates greater vegetation productivity (Santin-Janin et al., 2009) and, consequently, greater litter production. Our findings are likely attributed to the buildup of plant debris in the soil beneath plant canopy, resulting in the release of phosphorus when this organic material decomposes. The presence of organic substances may result in the liberation of organic acids (Shan et al., 2008), which can potentially diminish the capacity of soil particles to retain phosphorus (Dossa et al., 2008; Ge et al., 2019), thereby enhancing the availability of extractable phosphorus (Yao et al., 2019). Furthermore, owing to the deep root distribution of shrubs, phosphorus accumulation may occur in the deeper soil layers.
Our findings indicate that the fertile island effects in surface and subsurface soil depths are of similar magnitude across Tibetan Plateau drylands, at least under the shrub species that we assessed. However, different environmental factors regulated the fertile islands in surface and subsurface soil depths. The influence of climatic factors (precipitation of wettest quarter, precipitation seasonality, and temperature of driest quarter) on the fertile island effect was primarily evident in the soil surface layer. Increasing precipitation seasonality may strengthen the fertile islands effect of soil carbon in the surface layer, while climate warming may reduce this fertile island effect. In deeper soil layers, the fertile island effect is regulated by vegetation and soil texture, suggesting the importance of vegetation in redistributing nutrients in deeper soils. Our findings offer fresh perspectives on the variations in crucial factors influencing the fertility of islands at different depths, which is a significant characteristic observed in dryland ecosystems across the globe.