Figure 6 Piecewise SEM describing how biotic and abiotic variables
affect ecosystem multifunctionality. Numbers on the arrows represent all
the path coefficients and squared multiple correlations
(R 2) in the model. Solid lines indicate
significant paths and dashed lines represent paths that were not
significant. Positive path coefficients are pink and negative path
coefficients are blue. A single asterisk (*) and double asterisk (***)
indicate a significant difference between the variables at P < 0.05 and P < 0.001, respectively.
4 DISCUSSION
4.1 The diversity and composition of different trophic levels link to
ecosystem functions
Exploring the spatial distribution patterns and mechanisms of biological
species diversity on Earth has always been an important part of
ecological and biodiversity research. Our study found a correlation
between species diversity indices of different trophic levels,
especially in terms of species richness indices (Figure S4), which is
consistent with previous findings (Wang and Xing, 2021) indicating that
species diversity can be an important indicator reflecting community
dynamics. Meanwhile, plant communities undergo changes due to natural
disturbances in the form of succession, resulting in changes to the
community environment and effects on herbivores, pollinators and
microbes (Beck et al., 2015; Tao et al., 2023). The relationship between
plants and microbes can be understood from the mechanism of plant–soil
feedback, where different plant communities produce different types of
litter and root exudates, affecting soil organic carbon content and
thereby changing the composition and structure of soil microbial
communities (Smith et al., 2015). Researchers found that the influence
of plant diversity on multifunctionality was indirectly influenced by
shifts in soil biodiversity and plant cover (Delgado-Baquerizo et al.,
2020). This underscores the importance of incorporating soil
biodiversity into policy and management initiatives aimed at
safeguarding the functionality of terrestrial ecosystems on a global
scale. The activities of different types and quantities of rodents,
through digging and seed dispersal behaviors, have been shown to
influence the nutrient characteristics of the soil and plant species
(Pech et al., 2007; Wesche et al., 2007), leading to changes in the
composition of the surrounding plant community (Figure S6).
Ecosystem functions are closely related to the composition of keystone
species in biological communities. The realization of ecosystem
functions often depends on certain keystone species in the biological
community, such as important predators, pollinators, and major tree
species in forests (Raffard et al., 2021; Joshi et al., 2022). Our study
found that keystone species in plant, animal, and microbial communities
were interconnected with indicators of ecosystem functions (Figure S9),
suggesting that biodiversity influences ecosystem functions and
stability through interactions of species traits at multiple trophic
levels (Turnbull et al., 2016). At the same time, our results provided
evidence for the theory that ecological niche differentiation is
necessary to maintain species coexistence (MacArthur, 1970). For
example, the similarity of soil functional requirements of some plants
(e.g., Thalictrum aquilegiifolium var. sibiricum ,Sibbaldianthe bifurca , Saussurea pulchra , andEuphorbia altotibetica , Figure S9a) and their different
ecological niche widths in the community (Figure 1b) suggested that
there was obvious competition for resources in the plant community,
which affected the differences in the distribution of individual plants
in the community. It has been shown that species ecological niche
differentiation occurs during the construction of functional groups in
grasslands due to environmental factor constraints, resulting in the
coexistence of some species and the loss of others (Dray and Legendre,
2008), where environmental factors determine which traits are included
in functional groups that could be clustered in localized communities
(Ter Braak et al., 2012). Species ecological niche differentiation and
interspecific interactions based on functional traits thus play a
decisive role in plant community construction and diversity maintenance
(Kraft et al., 2008). In different vegetation areas, microbial key
species were also subject to limitations on ecosystem functioning
(Figures S5, S9b and S9c). The planctomycete_A-2 belongs to
Planctomycetes, which have the ability to utilize diverse carbon
compounds and can survive solely on high-molecular-weight dextran as
their carbon source, thereby participating in the carbon cycle (Boedeker
et al., 2017). Consequently, it was unsurprising to find that
planctomycete_A-2 was susceptible to nitrogen and carbon limitation in
our study. Meanwhile, Pseudomonas_frederiksbergensis has shown
potential to solubilize insoluble mineral phosphate (Zeng et al., 2016),
which promotes the release of effective phosphorus, and plant uptake of
phosphorus plays a significant role in promoting plant growth. We also
found that Pseudomonas_frederiksbergensis and the total
phosphorus content had a highly significant positive correlation.Xenodidymella_camporesii and Verrucaria_viridigrana belong to the phylum Ascomycota, which has been reported to tolerate
stressful conditions such as low nutrient availability, leading to more
efficient use of resources in challenging environments (Chen et al.,
2017). This could explain the lack of significant correlation with most
soil functions.
Rodents are the primary prey of many carnivores, while also consuming
plant seeds and fruits, thereby converting plant energy into animal
energy and dispersing plant seeds to different locations, promoting
biodiversity maintenance (Yu et al., 2023). Our research found that the
abundance of root voles was positively correlated with soil total
phosphorus content, nitrate nitrogen content, and aboveground biomass
(Figure S9d). A previous study suggested that the metabolic activities
of root voles produce urea and provide abundant nitrogen resources for
the areas surrounding their burrows, thus making the environment
suitable for plant survival and growth (Li et al., 2021). Plateau pikas,
as rodents mainly active underground, contribute to soil ventilation and
water infiltration through gnawing, digging, and excretion behaviors,
bringing organic matter and nutrients to the soil surface, enhancing
ecosystem functions, and providing favorable conditions for plant growth
(Zhang et al., 2003).
4.2 The pathways of multitrophic diversity drive ecosystem
multifunctionality
Most studies have begun to focus on changes in ecosystem
multifunctionality at different trophic levels. Our study showed that
multitrophic-level diversity had a greater impact on ecosystem
multifunctionality than the diversity of a single biome, suggesting that
maintaining ecosystem multifunctionality requires the joint
participation of biomes (Figures 4, 5 and 6), which is consistent with
the results of others (Schuldt et al., 2018; Luo et al., 2022; Mori et
al., 2023). Therefore, the relationship between biodiversity and
ecosystem multifunctionality is more dependent on diversity at the level
of multiple communities, and researchers should carry out their analyses
at large spatial scales rather than in single communities when
considering species diversity.
In addition to this, we found that ecosystem functions such as soil and
water conservation, soil fertility, nutrient cycling and transformation,
and community productivity were mediated by different single communities
in addition to multitrophic level diversity (Figure 4). Due to the
similarity in functional traits among different trophic groups and the
low redundancy between trophic levels, different trophic groups support
different functions (Soliveres et al., 2016; Luo et al., 2022). Plant
diversity had a positive effect on water conservation. The increase in
water conservation may result from increasing vegetation coverage, which
could lead to a reduction in soil evaporation and enhancement of
rainfall interception, as well as an increase in soil carbon and
nitrogen levels (Figure S10). This is because the presence of soil
organic matter can improve soil porosity, thereby boosting both soil
water infiltration and retention (Zhu et al., 2019; Zhao et al., 2023).
Plant community diversity, on the other hand, had a negative effect on
community productivity, which may have been due to the fact that
increased plant species richness increased competition among plants, and
plant competition has been shown to stimulate an increase in plant
litter and root secretions (Laganière et al., 2015). Our study in turn
confirmed that there were differences in resource utilization strategies
among different keystone species in the plant community, and therefore
community productivity can be highly unstable in this situation,
producing a downward trend. Soil microbes play an important role in
driving EMF (Jing et al., 2015; Delgado-Baquerizo et al., 2020). The
higher the soil bacterial diversity, the more important a role it plays
in nutrient cycling and transformation functions, and different
microbial species are strongly associated with different ecosystem
functions. Maintaining a higher richness of microbial taxa is crucial
for supporting increased functional redundancy and diversity, which
helps explain why greater biodiversity is necessary to sustain more
ecosystem functions (Wagg et al., 2019).
Here, we discovered that considering the characteristics of bacterial
communities often proved to be a more effective predictor of ecosystem
multifunctionality compared to considering fungal communities (Figure
4). This was due to the distribution of metabolic tasks among
microorganisms, creating synergies between microorganisms with distinct
physiological characteristics, such as those between fungi and bacteria.
This highlights the significance of interconnections between microbial
communities in influencing ecosystem functions, suggesting that these
hidden synergies may have a broader and more significant ecological
impact on soil microbial functions than previously recognized (Kohlmeier
et al., 2005; Deveau et al., 2018). Our study also found that soil
fertility, grassland productivity, and ecosystem multifunctionality
increased significantly with rodent community diversity, and that, in
general, mammals at low trophic levels, such as hares and rats, are
dependent on food resources and shelter provided by good ecological
niches. However, we also suspect that anthropogenic activities may be
replacing top-down ecological effects that are partially independent of
top predators. These ecological effects are not mediated by top
predators but directly affect small mammals (Smiley et al., 2020; She et
al., 2023).
Changes in the diversity of multiple trophic levels can be explained by
variations in the aboveground plant niche breadths, and as the niche
expands, the diversity of multiple trophic levels and ecosystem
multifunctionality also increase (Figure 6). The response of plant
species to interspecific competition involves altering their niches to
reduce overlap with other species, making them more complementary in
resource utilization (Eisenhauer et al., 2019). Plant communities with
high diversity select for enhanced niche differentiation among species,
reducing interspecific competition without increasing intraspecific
competition. This mechanism may strengthen the relationship between
biodiversity and ecosystem functioning (Amyntas et al., 2023). The
relationship between plant diversity and ecosystem functioning depends
not only on interactions between plants but also on interactions within
and between different trophic levels of the food chain (Barnes et al.,
2020; Albert et al., 2022). This emphasizes the importance of plants as
primary producers in connecting aboveground consumers and belowground
decomposers, thus influencing changes in multi-trophic level biological
communities.
4.3 Altitude and pH influence ecosystem multifunctionality
Biodiversity is not the sole or primary driving factor of ecosystem
multifunctionality (Giling et al., 2019); climate and abiotic factors
also drive ecosystem functions (Grytnes and McCain, 2013). We found that
biotic factor of altitude and pH directly explained changes in ecosystem
multifunctionality (Figures 5 and 6), which aligned with previous
evidence (Hu et al., 2020; Luo et al., 2022). Altitude serves as a
predictive factor for patterns of species diversity and community
composition under environmental changes. With increasing altitude,
temperature decreases, solar radiation increases, and wind strength
intensifies, putting significant stress on the physiological and
survival strategies of species (Galván-Cisneros et al., 2023). Studies
had shown that ecosystem multifunctionality significantly decreases with
increasing altitude (Chen et al., 2022),which is contrary to our
results. The reason for this discrepancy may be that the relationship
between altitude and ecosystem multifunctionality is not simply linear
but follows a single-peak curve. Research on the dependence of ecosystem
multifunctionality on altitude in the Qinghai-Tibet Plateau indicated
that there was a critical point around 3900 m where the relationship
between ecosystem multifunctionality and altitude changes (Wang et al.,
2023b). Our study sites were located near this critical point, which
might explain why there was a positive correlation between altitude and
ecosystem multifunctionality in our findings. Soil pH directly affects
processes such as mineral weathering, organic matter mineralization, and
humification in soils, which have a significant impact on the status of
nutrient ions in soils. Most studies have confirmed that soil
acidification reduces ecosystem functionality (Delgado-Baquerizo et al.,
2016; Wei et al., 2022),which is inconsistent with our research
findings. Through our study of sample points, we found that
high-altitude grasslands with lower pH and higher ecosystem
multifunctionality show obvious shrub encroachment. The pH content of
shrub-encroached high-altitude grasslands tends to be acidic (Ma et al.,
2022),and shrub encroachment can enhance ecosystem functionality,
specifically in terms of carbon sequestration, soil fertility, and
hydrological functions (Ding and Eldridge, 2023). These results support
our conclusion.
5 CONCLUSIONS
In summary, our results indicate that key species at different trophic
levels are interconnected with ecosystem functions, leading to species
niche differentiation within communities and influencing the formation
of biodiversity. The interconnections of diversity at a single trophic
level, especially richness indices, highlight the importance of
biodiversity in maintaining food web structures and functions within
biological communities. The greater the number of species in an
ecosystem, the more diverse the food base supply, ensuring ecosystem
stability. Furthermore, we confirmed the role of multi-trophic
diversity, namely biodiversity (i.e., plant diversity, bacterial
diversity, and rodent diversity), in maintaining ecosystem functions,
further emphasizing the critical importance of conserving biodiversity
in sustaining ecosystem functions. Lastly, abiotic factors such as
altitude alter ecosystem multifunctionality by affecting soil
environments, and multitrophic diversity directly impacts ecosystem
multifunctionality. Therefore, in the face of increasing human
activities and climate change, the maintenance of multifunctionality
through multitrophic diversity is crucial and requires our utmost
attention.
AUTHOR CONTRIBUTIONS
All authors contributed to the article. H.S., H.Z., Z.W., and H.L., made
efforts to define the topic of the study and guided the writing of the
article. The help of R.Q. and T.C. solved the problem of rodent capture.
L.M., Z.Z., X.H., and F.Y. solved the problems encountered during data
processing and graphing. J.Y., X.L., S.L., J.W., H.A. and Z.S.
participated in the field survey and sampling work.
DATA AVAILABILITY
Data will be made available on request.
ACKNOWLEDGMENTS
This study was funded by the National Natural Science Foundation of
China (32371684), the National Natural Science Foundation of China Joint
Fund Project (U21A20186, U20A2006), the Second Tibetan Plateau
Scientific Expedition and Research (STEP) Program (2019QZKK0302–02),
and the International Cooperation Project of Key Research and
Development and Transformation in Qinghai Province (2024–HZ–810).
CONFLICTS OF INTEREST
The authors declare no competing interests.
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