Plant diversity increases soil microbial activity and soil carbon storage
Open accessAbstract
Plant diversity strongly influences ecosystem functions and services, such as soil carbon storage. However, the mechanisms underlying the positive plant diversity effects on soil carbon storage are poorly understood. We explored this relationship using long-term data from a grassland biodiversity experiment (The Jena Experiment) and radiocarbon (14C) modelling. Here we show that higher plant diversity increases rhizosphere carbon inputs into the microbial community resulting in both increased microbial activity and carbon storage. Increases in soil carbon were related to the enhanced accumulation of recently fixed carbon in high-diversity plots, while plant diversity had less pronounced effects on the decomposition rate of existing carbon. The present study shows that elevated carbon storage at high plant diversity is a direct function of the soil microbial community, indicating that the increase in carbon storage is mainly limited by the integration of new carbon into soil and less by the decomposition of existing soil carbon. The mechanisms driving soil carbon storage, one of the largest stores of terrestrial carbon, remain poorly understood. Here, the authors present data from the long-term Jena Experiment on grassland biodiversity, showing that elevated carbon storage at high plant diversity is a direct function of increased soil microbial activity.
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Written from the paper’s full text.
TL;DR
Plant diversity in a long-term grassland experiment increased soil carbon storage mainly by enhancing root carbon inputs and microbial activity, which promoted accumulation of new carbon into slow-cycling soil organic matter. Radiocarbon modelling and path analysis indicated that decomposition/priming of old carbon was less important than the integration of new carbon.
Key findings
- Nine years after establishment, topsoil (0–5 cm) soil carbon concentration had increased by 27% to 4.74 ± 0.28 g kg⁻¹ and was highly correlated with sown plant species richness (P < 0.001); legume presence negatively affected soil carbon concentration (P = 0.040).
- Path analysis found direct positive effects of plant species richness on root carbon inputs and microbial basal respiration, but no direct relationship between plant species richness or legumes and soil carbon storage; increased soil carbon storage was directly related to higher microbial activity (best model: χ²₄ = 5.35, P = 0.253; RMSEA = 0.065).
- Radiocarbon-based modelling showed high-diversity plots (richness 8, 16, 60) accumulated 21.8% more carbon than low-diversity plots (richness 1, 2, 4), while high-diversity plots primed on average only 9.6% of old carbon; new carbon sequestration and old carbon loss were positively correlated (R² = 0.863, P = 0.007).
- In a complementary ¹³CO₂ labelling experiment, label uptake into root-associated microbial PLFAs increased with plant species richness (F₁,₉ = 6.99, P = 0.027), and bacterial and fungal diversity increased with plant species richness (ANOVA PSR F-values: fungi 12.06*, bacteria 19.37*).
Methods
- The study used The Jena Experiment, a grassland biodiversity experiment with 81 plots (one monoculture abandoned from 82 planned), sown species richness levels of 1, 2, 4, 8, 16 and 60 species and 1–4 plant functional groups; main analyses focused on 0–5 cm topsoil, with some 0–30 cm data.
- Analyses included type I sum-of-squares ANOVA, path analysis using maximum likelihood in lavaan (N = 81), and a three-pool radiocarbon model (fast, slow, passive pools) constrained by soil carbon stocks and ¹⁴C, implemented in SoilR.
- Additional measurements included basal respiration, fine root biomass/carbon, microbial biomass by chloroform fumigation, TRFLP for bacterial and fungal diversity, and an Ecotron ¹³CO₂ labelling experiment with PLFA analysis.
Limitations
- The paper states that it cannot rule out that the proposed mechanisms are particularly pronounced at The Jena Experiment field site; root exudation could not be determined within the Jena Experiment, so an Ecotron experiment was used instead.
- The paper states that no basal respiration data were available for arable control plots needed for model spin-up, so microbial biomass was used instead; the real carbon input and the extent of the transfer coefficient could not be evaluated, so a set of different models was applied.
- Evident from the text: the main mechanistic analyses are based on the top 0–5 cm, and generalization to deeper layers is inferential; high-diversity treatments had uneven replication, with only 4 plots at richness 60, which may limit robustness of diversity-level estimates.
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Claims the paper makes
0 of 39 cited sources checked · 0 supported · 0 flagged
- Findingown data§ Results · p. 2
The increase in soil carbon concentration was highly correlated with sown plant species richness.
“The increase in soil carbon concentration was highly correlated with sown plant species richness ( P o 0.001, Fig. 1).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
Increased soil carbon storage was only directly related to higher microbial activity.
“Importantly, the increased soil carbon storage was only directly related to higher microbial activity.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
The presence of legumes negatively affected soil carbon concentration.
“The presence of legumes negatively affected soil carbon concentration ( P ¼ 0.040, Fig. 1)”
Quote in the paperNot a substantial claim - Findingown data§ Results · p. 2
In 2011, nine years after establishment of The Jena Experiment, soil carbon concentration in the topsoil layer (0-5 cm, averaged across all diversity treatments) had increased by 27% to 4.74 ± 0.28 g kg−1.
“In 2011, 9 years after the establishment of The Jena Experiment, soil carbon concentration in the topsoil layer (0-5 cm, averaged across all diversity treatments) had increased by 27% to 4.74 ± 0.28 g kg 1 (mean ± s.e.m.).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
There was no direct relationship between plant species richness and soil carbon storage.
“There was no direct relationship between plant species richness or the presence of legumes and soil carbon storage”
Quote in the paperSubstantial claim
- Findingown data§ Results · p. 2
Plant species richness had direct positive effects on metabolic activity of soil microorganisms, measured as basal respiration.
“Path analyses revealed that plant species richness had direct positive effects on root carbon inputs as well as metabolic activity of soil microorganisms, measured as basal respiration (Fig. 3a,b; Methods).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
There was no direct relationship between the presence of legumes and soil carbon storage.
“There was no direct relationship between plant species richness or the presence of legumes and soil carbon storage”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
Plant species richness had direct positive effects on root carbon inputs.
“Path analyses revealed that plant species richness had direct positive effects on root carbon inputs as well as metabolic activity of soil microorganisms, measured as basal respiration (Fig. 3a,b; Methods).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
Radiocarbon-based modelling of SOM storage suggested that increased carbon storage in plots with higher plant species richness was mainly due to accumulation of 'new' carbon.
“Radiocarbon-based modelling of SOM storage (Methods) suggested that the increased carbon storage in plots with higher plant species richness was mainly due to the accumulation of 'new' carbon (Fig. 4a).”
Quote in the paperSubstantial claim - Findingown data§ Discussion · p. 3
Carbon uptake in rhizospheric microorganisms at high plant diversity was increased compared with low plant diversity.
“Indeed, carbon uptake in rhizospheric microorganisms at high plant diversity was increased compared with low plant diversity (Fig. 5) as indicated by a complementary 13 CO2 labelling experiment in the Ecotron facility in Montpellier (Methods).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
High-diversity plots primed on average only 9.6% of 'old' carbon.
“At the same time, high-diversity plots primed on average only 9.6% of 'old' carbon.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
High-diversity plots (richness levels 8, 16 and 60) accumulated 21.8% more carbon compared with low-diversity plots (richness levels 1, 2 and 4).
“Highdiversity plots (richness levels 8, 16 and 60) accumulated 21.8% more carbon compared with low-diversity plots (richness levels 1, 2 and 4).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Carbon losses were mainly observed in the fast-cycling carbon.
“Interestingly, carbon losses were mainly observed in the fast-cycling carbon (Fig. 4c)”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 2
The potential chain of causality from plant species richness through root carbon inputs and microbial activity to soil carbon storage was the best model that adequately explained the data.
“This potential chain of causality was the best model that adequately explained the data (Supplementary Table 2; Supplementary Notes 1-4)”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Findingcitation§ Discussion · p. 4
The positive plant diversity effect on soil carbon storage, root biomass and soil microbial biomass was also found for deeper soil layers in The Jena Experiment.
“In contrast, the positive plant diversity effect on soil carbon storage (Table 1), root biomass 39 and soil microbial biomass 40 was also found for deeper soil layers in The Jena Experiment.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [39] Ravenek, J. M. et al. Long-term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time. Oikos 123, 1528–1536 (2014) . source
Not checked in the free run.
- cites [40] Guenay, Y., Ebeling, A., Steinauer, K., Weisser, W. W. & Eisenhauer, N. Transgressive overyielding of soil microbial biomass in a grassland plant diversity gradient. Soil Biol. Biochem. 60, 122–124 (2013) . source
Not checked in the free run.
- Findingown data§ Discussion · p. 3
Label enrichment in roots and root sugars was not affected by plant diversity.
“the label enrichment in roots and root sugars was not affected by diversity (data not shown).”
Quote in the paperSubstantial claim - Findingown data§ Discussion · p. 3
Bacterial and fungal diversity increased with higher plant species richness in The Jena Experiment.
“In fact, bacterial and fungal diversity increased with higher plant species richness in The Jena Experiment (Table 1; Fig. 6).”
Quote in the paperNot a substantial claim - Findingown data§ Methods · p. 7
Basal respiration and microbial biomass were highly correlated.
“Basal respiration and microbial biomass were highly correlated (Fig. 3c).”
Quote in the paperSubstantial claim - Findingown data§ Discussion · p. 4
The increase in total nitrogen content was highly correlated with the increase in soil organic carbon.
“Although the increase in total nitrogen content was highly correlated with the increase in soil organic carbon ( R 2 ¼ 0.79, P o 0.001)”
Quote in the paperNot a substantial claim - Findingown data§ Discussion · p. 3
There was slightly but significantly more microbial carbon per unit root carbon in high-diversity plots.
“indicated by slightly but significantly more microbial carbon per unit root carbon in high-diversity plots ( R 2 ¼ 0.06, P ¼ 0.025).”
Quote in the paperSubstantial claim - Findingown data§ Methods · p. 7
Implementing the transfer coefficient between the fast and the slow pool depending on soil microbial biomass resulted in good agreement between model predictions and measured data.
“Implementing the transfer coefficient between the fast and the slow pool depending on soil microbial biomass resulted in good agreement between model predictions and measured data (Table 3; Supplementary Table 3; and Supplementary Fig. 3).”
Quote in the paperSubstantial claim - Findingown data§ Methods · p. 6
Fine root biomass was highly correlated to the total root biomass.
“and was highly correlated to the total root biomass ( R 2 ¼ 0.66, P o 0.001).”
Quote in the paperNot a substantial claim - Interpretationown data§ Results · p. 2
Fine root carbon inputs and microbial activity are the main drivers explaining the positive effect of plant diversity on soil carbon storage.
“showing that fine root carbon inputs and microbial activity are the main drivers explaining the positive effect of plant diversity on soil carbon storage.”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Interpretationown data§ Results · p. 3
The transfer or processing rate of carbon from fast- to slow-cycling carbon is proportional to soil microbial biomass and activity.
“(conversion efficiency, Methods) of carbon from fast- to slowcycling carbon, which is-in accordance with our path modelproportional to soil microbial biomass and activity (Fig. 3c).”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Interpretationcitation§ Discussion · p. 4
Lower root biomass on legume plots likely caused the negative legume effect on soil carbon storage.
“the lower root biomass on legume plots 29 likely caused the negative legume effect on soil carbon storage.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [29] Bessler, H. et al. Aboveground overyielding in grassland mixtures is associated with reduced biomass partitioning to belowground organs. Ecology 90, 1520–1530 (2009) . source
Not checked in the free run.
- Interpretationreasoning§ Discussion · p. 3
Higher microbial label uptake was most likely attributed to higher levels of root exudation with increasing plant diversity.
“This showed higher microbial label uptake was most likely attributed to higher levels of root exudation with increasing plant diversity, as the soil microbial community was sampled immediately after a 3-week label application and the label enrichment in roots and root sugars was not affected by diversity (data not shown).”
Quote in the paperSubstantial claim- Rests on an assumption: The 13CO2 labelling experiment in the Ecotron facility is representative of root exudation effects on rhizospheric microorganisms in The Jena Experiment field plots.
- Interpretationreasoning§ Discussion · p. 4
The activity and composition of soil microbial communities can serve as a proxy for carbon transfer into sustainable slow-cycling forms of soil carbon.
“Thus, our results suggest that the activity and composition of soil microbial communities can serve as a proxy for carbon transfer into sustainable slow-cycling forms of soil carbon”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Interpretationreasoning§ Discussion · p. 4
Higher levels of carbon inputs to the soil and more favourable microclimatic conditions linked to more diverse plant communities result in more active, more abundant and more diverse soil microbial communities.
“In summary, higher levels of carbon inputs to the soil and more favourable microclimatic conditions linked to more diverse plant communities result in more active, more abundant and more diverse soil microbial communities.”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Premisecitation§ Discussion · p. 3
Increasing diversity of soil microbial communities increases microbial respiration.
“Third, increasing diversity of soil microbial communities increases microbial respiration 34 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [34] Bell, T., Newman, J. A., Silverman, B. W., Turner, S. L. & Lilley, A. K. The contribution of species richness and composition to bacterial services. Nature 436, 1157–1160 (2005) . source
Not checked in the free run.
- Interpretationcitation§ Discussion · p. 4
The plant diversity effect may be strongest in the topsoil and decrease with soil depth.
“Despite the consistency of the plant diversity effect, it may be strongest in the topsoil and decrease with soil depth 25,39 , which may be due to the fact that root carbon inputs 24,39 and microbial activity 41 decrease with soil depth.”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [25] Steinbeiss, S. et al. Plant diversity positively affects short-term soil carbon storage in experimental grasslands. Glob. Change Biol. 14, 2937–2949 (2008) . source
Not checked in the free run.
- cites [39] Ravenek, J. M. et al. Long-term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time. Oikos 123, 1528–1536 (2014) . source
Not checked in the free run.
- Interpretationcitation§ Discussion · p. 4
Microbial products associated with increased microbial respiration such as microbial necromass end up in slow-cycling SOM pools in the form of reduced organic material.
“Thus, microbial products associated with increased microbial respiration such as microbial necromass end up in slow-cycling SOM pools in the form of reduced organic material 10,14 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [10] Schmidt, M. W. I. et al. Persistence of soil organic matter as an ecosystem property. Nature 478, 49–56 (2011) . source
Not checked in the free run.
- cites [14] Miltner, A., Bombach, P., Schmidt-Bruecken, B. & Kaestner, M. SOM genesis: microbial biomass as a significant source. Biogeochemistry 111, 41–55 (2012) . source
Not checked in the free run.
- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Interpretationreasoning§ Discussion · p. 4
Microbial activity increases the turnover rates of root litter and exudates as indicated by increased microbial respiration.
“Microbial activity increases the turnover rates of root litter and exudates as indicated by increased microbial respiration.”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
- Interpretationreasoning§ Discussion · p. 4
Plant diversity and associated soil microbial communities can significantly contribute to sequestration of atmospheric carbon dioxide.
“and that plant diversity and associated soil microbial communities can significantly contribute to sequestration of atmospheric carbon dioxide.”
Quote in the paperSubstantial claim - Premisecitation§ Methods · p. 6
Plant species richness was not closely linked to basal respiration and root standing biomass in the first years of the experiment.
“Eisenhauer et al. 20 and Ravenek et al. 39 showed that plant species richness was not closely linked to basal respiration and root standing biomass in the first years of the experiment; therefore, we did not use data from the first 2 years in the present analyses.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Eisenhauer et al. 20
Not checked in the free run.
- cites Ravenek et al. 39
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- Interpretationown data§ Discussion · p. 3
A shift in the metabolic activity of soil microorganisms towards anabolic activity with plant diversity resulted in increased necromass accumulation over time.
“And finally, a shift in the metabolic activity of soil microorganisms towards anabolic activity with plant diversity resulting in increased necromass accumulation over time 23 , indicated by slightly but significantly more microbial carbon per unit root carbon in high-diversity plots ( R 2 ¼ 0.06, P ¼ 0.025).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [23] Liang, C., Cheng, G., Wixon, D. L. & Balser, T. C. An Absorbing Markov Chain approach to understanding the microbial role in soil carbon stabilization. Biogeochemistry 106, 303–309 (2011) . source
Not checked in the free run.
- Premisecitation§ Methods · p. 6
Particular specific monounsaturated fatty acids such as PLFA 16:1ω5, 16:1ω7, 17:1ω8 and 18:1ω7 are highly enriched by root-derived organic carbon in label experiments.
“Particular specific monounsaturated fatty acids such as PLFA 16:1 o 5, 16:1 o 7, 17:1 o 8 and 18:1 o 7 were shown to be highly enriched by root-derived organic carbon in label experiments 54,55 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [54] Esperschuetz, J. et al. Microbial response to exudates in the rhizosphere of young beech trees (Fagus sylvatica L.) after dormancy. Soil Biol. Biochem. 41, 1976–1985 (2009) . source
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- cites [55] Treonis, A. M. et al. Identification of groups of metabolically-active rhizosphere microorganisms by stable isotope probing of PLFAs. Soil Biol. Biochem. 36, 533–537 (2004) . source
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- Study design and conductcitation§ Methods · p. 6
The twelve monoliths reflected the diversity gradient of The Jena Experiment on a smaller scale.
“These 12 monoliths reflected the diversity gradient of The Jena Experiment on a smaller scale 51 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [51] Milcu, A. et al. Functional diversity of leaf nitrogen concentrations drives grassland carbon fluxes. Ecol. Lett. 17, 435–444 (2014) . source
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- Study design and conductnone given§ Methods · p. 6
Within the experimental setup of The Jena Experiment it is not possible to determine the effects of plant diversity on the amount of root exudation.
“Within the experimental setup of The Jena Experiment it is not possible to determine the effects of plant diversity on the amount of root exudation.”
Quote in the paperSubstantial claim - Premiseown data§ Methods · p. 7
Including microbial biomass as an additional pool in the dynamical system leads to unrealistic nonlinear behaviour not supported by the authors' own observations.
“Notice that we do not represent the microbial biomass as an additional pool in our dynamical system because it leads to unrealistic nonlinear behaviour 64 not supported by our own observations.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [64] Wang, Y. P. et al. Oscillatory behavior of two nonlinear microbial models of soil carbon decomposition. Biogeosci. Discuss. 10, 19661–19700 (2013) . source
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- May combine two claims. 18
- Classified as probably not a substantial claim. 17
May combine two claims.
Findingown data§ Results · p. 2Other plant functional groups or their richness did not significantly influence soil carbon concentration.
“while other plant functional groups or their richness did not significantly influence carbon concentration (Table 1; Supplementary Table 1).”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown data§ Results · p. 3Sequestration of 'new' carbon and loss of existing 'old' carbon depended on plant species richness.
“and depended on plant species richness.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ Results · p. 3In the slow-cycling carbon, losses were reduced at higher plant diversity and more 'new' carbon was stored.
“whereas in the more sustainable slow-cycling carbon these losses were reduced at higher plant diversity, and more 'new' carbon was stored in the slow-cycling carbon (Fig. 4d).”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Findingown data§ Methods · p. 6Fine root biomass comprised on average 80.9% (±16.8 s.d.) of the total root biomass.
“Fine root biomass comprised on average 80.9% ( ± 16.8 s.d.) of the total root biomass”
Quote in the paperNot a substantial claimMay combine two claims.
Findingcitation§ Discussion · p. 4The presence of legumes did not increase root biomass production and the metabolic activity of soil microorganisms in The Jena Experiment.
“However, the presence of legumes did not increase root biomass production and the metabolic activity of soil microorganisms in The Jena Experiment 20,38 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [20] Eisenhauer, N. et al. Plant diversity effects on soil microorganisms support the singular hypothesis. Ecology 91, 485–496 (2010) . source
Not checked in the free run.
- cites [38] Bessler, H. et al. Nitrogen uptake by grassland communities: contribution of N-2 fixation, facilitation, complementarity, and species dominance. Plant Soil 358, 301–322 (2012) . source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Findingnone given§ Methods · p. 7After establishment of experimental plots, the fine root biomass distribution was similar to meadows, with the main amount of roots in the topsoil and strongly decreasing towards deeper depths.
“In contrast, after establishment of experimental plots, the fine root biomass distribution was similar to meadows with the main amount of roots in the topsoil, strongly decreasing towards deeper depths.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationown data§ Discussion · p. 3Storage of soil carbon is governed by the metabolic activity of soil microbes, which is mediated by plant diversity via higher root inputs and other yet unidentified mechanisms.
“The results reported in this study indicate that storage of soil carbon is governed by the metabolic activity of soil microbes, which is mediated by plant diversity via higher root inputs and other yet unidentified mechanisms, as suggested by the direct relation in the path model (Fig. 3a).”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
May combine two claims.
Interpretationreasoning§ Results · p. 3Slow-cycling carbon (turnover time = 47.7 years) dominates the longer-term dynamics of the total stock, whereas priming and other effects of the fast-cycling carbon (turnover time = 4.0 years) become less important over longer time scales.
“Consequently, the slow-cycling carbon (turnover time ¼ 47.7 years, Supplementary Table 3) dominates the longer-term dynamics of the total stock, whereas priming and other effects of the fast-cycling carbon (turnover time ¼ 4.0 years) become less important over longer time scales.”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
May combine two claims.
Premisecitation§ Discussion · p. 3A denser vegetation in highly diverse plant communities reduces evaporation from the topsoil, which in turn promotes higher soil microbial activity and growth.
“First, a denser vegetation in highly diverse plant communities reduces evaporation from the topsoil, which in turn promotes higher soil microbial activity and growth 30 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [30] Lange, M. et al. Biotic and Abiotic properties mediating plant diversity effects on soil microbial communities in an experimental grassland. PLoS ONE 9, e96182 (2014) . source
Not checked in the free run.
May combine two claims.
Premisecitation§ Discussion · p. 3Soil carbon storage is linked to root inputs including root exudation, which is known to change the activity and composition of the microbial community.
“Second, soil carbon storage is linked to root inputs 31,32 including root exudation, which is known to change the activity and composition of the microbial community 33 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [31] Kramer, C. et al. Recent (< 4 year old) leaf litter is not a major source of microbial carbon in a temperate forest mineral soil. Soil Biol. Biochem. 42, 1028–1037 (2010) . source
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- cites [32] Rasse, D. P., Rumpel, C. & Dignac, M. F. Is soil carbon mostly root carbon? Mechanisms for a specific stabilisation. Plant Soil 269, 341–356 (2005) . source
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- cites [33] Bais, H. P., Weir, T. L., Perry, L. G., Gilroy, S. & Vivanco, J. M. The role of root exudates in rhizosphere interations with plants and other organisms. Annu. Rev. Plant Biol. 57, 233–266 (2006) . source
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May combine two claims.
Interpretationreasoning§ Discussion · p. 4The mechanism proposed for the topsoil is very likely to also be relevant for deeper soil layers, but the decreased biological activity at the deeper soil layers has to be taken into account.
“Therefore, the mechanism proposed for the topsoil is very likely to also be relevant for deeper soil layers, but the decreased biological activity at the deeper soil layers has to be taken into account.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationcitation§ Discussion · p. 4The underlying relationships between plant diversity and soil carbon storage, soil microbial biomass and activity, and root biomass have been reported in several independent experimental settings, providing some evidence that the proposed links are of general significance.
“Although we cannot rule out that the proposed mechanisms are particularly pronounced at the field site of The Jena Experiment, the underlying relationships between plant diversity and soil carbon storage 17,24 , soil microbial biomass and activity 37,45,46 , and root biomass 24,47 have been reported in several independent experimental settings providing some evidence that the proposed links are of”
Quote in the paperSubstantial claim0 of 6 sources checked- cites [17] De Deyn, G. B. et al. Additional carbon sequestration benefits of grassland diversity restoration. J. Appl. Ecol. 48, 600–608 (2011) . source
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- cites [24] Fornara, D. A. & Tilman, D. Plant functional composition influences rates of soil carbon and nitrogen accumulation. J. Ecol. 96, 314–322 (2008) . source
Not checked in the free run.
- cites [37] Zak, D. R., Holmes, W. E., White, D. C., Aaron, D. P. & Tilman, D. Plant diversity, soil microbial communities, and ecosystem function: are there any links? Ecology 84, 2042–2050 (2003) . source
Not checked in the free run.
- cites [45] Eisenhauer, N. et al. Plant diversity effects on soil food webs are stronger than those of elevated CO2 and N deposition in a long-term grassland experiment. Proc. Natl Acad. Sci. USA 110, 6889–6894 (2013) . source
Not checked in the free run.
- cites [46] Spehn, E. M., Joshi, J., Schmid, B., Alphei, J. & Korner, C. Plant diversity effects on soil heterotrophic activity in experimental grassland ecosystems. Plant Soil 224, 217–230 (2000) . source
Not checked in the free run.
- cites [47] Mueller, K. E., Tilman, D., Fornara, D. A. & Hobbie, S. E. Root depth distribution and the diversity-productivity relationship in a long-term grassland experiment. Ecology 94, 787–793 (2013) . source
Not checked in the free run.
May combine two claims.
Interpretationreasoning§ Discussion · p. 4The positive plant diversity effect on carbon storage is driven by root carbon inputs, but mediated by the soil microbial community.
“The positive plant diversity effect on carbon storage is driven by root carbon inputs, but mediated by the soil microbial community.”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
May combine two claims.
Interpretationown data§ Discussion · p. 4The legume effect might be most pronounced in the topsoil as it was not significant when considering the top 30 cm of the soil.
“However, the legume effect might be most pronounced in the topsoil as it was not significant when considering the top 30 cm of the soil (Table 1).”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim. May combine two claims.
Study design and conductreasoning§ Methods · p. 6Statistical analyses were done with carbon concentrations and stocks were estimated using the mean of bulk soil density across all years in order to exclude physical aspects of bulk density.
“In order to exclude physical aspects of bulk density (for example, inter-annual differences caused by degree of winter freezing), statistical analyses were done with carbon concentrations (g kg 1 ) and stocks were estimated using the mean of bulk soil density across all years.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductreasoning§ Methods · p. 5Plots are arranged in four blocks to account for changes in abiotic factors with distance from the river.
“Plots are arranged in four blocks to account for changes in abiotic factors with distance from the river.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductcitation§ Methods · p. 5Initial physico-chemical properties varied across the field site and are considered in the block design of The Jena Experiment.
“The initial physico-chemical properties, such as pH (7.1-8.4), soil organic carbon (5-33 g C kg 1 ), and soil nitrogen concentrations (1.0-2.7 g N kg 1 ), varied across the field site and are considered in the block design of The Jena Experiment 26 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [26] Roscher, C. et al. The role of biodiversity for element cycling and trophic interactions: an experimental approach in a grassland community. Basic Appl. Ecol. 5, 107–121 (2004) . source
Not checked in the free run.
May combine two claims.
Interpretationown data§ Discussion · p. 3The missing direct path from legumes to microbial activity indicates minor importance of improvement of soil nitrogen status for soil microbial activity, and thus for carbon storage.
“The missing direct path from legumes to microbial activity indicates minor importance of improvement of soil nitrogen status (measured as changes in 15 N of bulk soil, Table 1) for soil microbial activity, and thus for carbon storage.”
Quote in the paperSubstantial claim- Rests on an assumption: The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
May combine two claims.
Premisecitation§ Discussion · p. 4Root carbon inputs and microbial activity decrease with soil depth.
“which may be due to the fact that root carbon inputs 24,39 and microbial activity 41 decrease with soil depth.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [24] Fornara, D. A. & Tilman, D. Plant functional composition influences rates of soil carbon and nitrogen accumulation. J. Ecol. 96, 314–322 (2008) . source
Not checked in the free run.
- cites [39] Ravenek, J. M. et al. Long-term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time. Oikos 123, 1528–1536 (2014) . source
Not checked in the free run.
- cites [41] Kramer, C. & Gleixner, G. Soil organic matter in soil depth profiles: distinct carbon preferences of microbial groups during carbon transformation. Soil Biol. Biochem. 40, 425–433 (2008) . source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ Methods · p. 7The ratio of the assumed fine root inputs between experimental plots and arable land is similar to the ratio of carbon input on meadows and arable lands reported in ref. 68.
“The ratio of the assumed fine root inputs between experimental plots and arable land is similar to the ratio of carbon input on meadows and arable lands reported in ref. 68”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [68] Schulze, E. D. et al. Importance of methane and nitrous oxide for Europe's terrestrial greenhouse-gas balance. Nat. Geosci. 2, 842–850 (2009) . source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductcitation§ Methods · p. 6The block effect, mainly accounting for differences in soil texture among the blocks, was included as a random factor and fitted first, followed by plant species richness and functional groups.
“The 'block effect', mainly accounting for differences in soil texture among the blocks 26 , was included as random factor and fitted first, followed by plant species richness and functional groups.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [26] Roscher, C. et al. The role of biodiversity for element cycling and trophic interactions: an experimental approach in a grassland community. Basic Appl. Ecol. 5, 107–121 (2004) . source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone given§ Methods · p. 6One monoculture plot was abandoned due to poor establishment, making the total number of plots 81 instead of the 82 originally set up.
“*One monoculture plot was abandoned due to poor establishment, making the total number of plots 81 instead of the 82, which were originally set up.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Interpretationown data§ Methods · p. 7On the basis of the path analysis findings, basal respiration was implemented as a proxy for the transfer between the fast and slow carbon pool.
“On the basis of our findings in the path analysis, we implemented a measure of soil microbial community activity (basal respiration) as a proxy for the transfer between the fast and slow carbon pool.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim. May combine two claims.
Premisecitation§ Methods · p. 6Fine root carbon is a measure of carbon input and basal respiration is a measure of microbial metabolic activity, and both were included as variables potentially mediating plant community effects.
“As variables potentially mediating plant community effects we included fine root carbon as measure of carbon input 31,32 and basal respiration as measure of microbial metabolic activity 20,34,37 .”
Quote in the paperSubstantial claim0 of 5 sources checked- cites [31] Kramer, C. et al. Recent (< 4 year old) leaf litter is not a major source of microbial carbon in a temperate forest mineral soil. Soil Biol. Biochem. 42, 1028–1037 (2010) . source
Not checked in the free run.
- cites [32] Rasse, D. P., Rumpel, C. & Dignac, M. F. Is soil carbon mostly root carbon? Mechanisms for a specific stabilisation. Plant Soil 269, 341–356 (2005) . source
Not checked in the free run.
- cites [20] Eisenhauer, N. et al. Plant diversity effects on soil microorganisms support the singular hypothesis. Ecology 91, 485–496 (2010) . source
Not checked in the free run.
- cites [34] Bell, T., Newman, J. A., Silverman, B. W., Turner, S. L. & Lilley, A. K. The contribution of species richness and composition to bacterial services. Nature 436, 1157–1160 (2005) . source
Not checked in the free run.
- cites [37] Zak, D. R., Holmes, W. E., White, D. C., Aaron, D. P. & Tilman, D. Plant diversity, soil microbial communities, and ecosystem function: are there any links? Ecology 84, 2042–2050 (2003) . source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ Methods · p. 7Low 14C values are common for arable fields in Europe.
“As described by other authors 66 , such low 14 C values are common for arable fields in Europe.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [66] Brodowski, S., Amelung, W., Haumaier, L. & Zech, W. Black carbon contribution to stable humus in German arable soils. Geoderma 139, 220–228 (2007) . source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Study design and conductnone given§ Methods · p. 7The value of the transfer coefficient is within the expected range.
“This value of the transfer coefficient is within the expected range.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductnone given§ Methods · p. 7The real carbon input and the extent of the transfer coefficient could not be evaluated.
“However, we could neither evaluate the real carbon input (carbon of standing fine root biomass and root exudates, root tips and so on) nor the extent of the transfer coefficient.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Study design and conductnone given§ Methods · p. 6For all models, the adequacy of the hypothesized structural relationships and the data was verified using χ2-tests and the residual mean square error of approximation.
“For all models, the adequacy of the hypothesized structural relationships and the data was verified using w 2 -tests and the residual mean square error of approximation.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown data§ Methods · p. 6In the final step, the minimal adequate (most parsimonious) model was identified based on the Bayes Information Criterion.
“In the final step, we identified the minimal adequate (most parsimonious) model based on the Bayes Information Criterion.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Interpretationcitation§ Methods · p. 7The passive pool is mainly needed to account for the low 14C values in the bulk soil organic matter.
“The passive pool is mainly needed to account for the low 14 C values in the bulk soil organic matter.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Study design and conductown data§ Methods · p. 7No data on basal respiration were available for arable control plots, which would have been needed for the spin up.
“Unfortunately, no data on basal respiration were available for arable control plots, which would have been needed for the spin up (please see section 'Spin up and parameter estimation for the spin up').”
Quote in the paperSubstantial claim
The paper relies on this without support.
The observed increase in soil organic matter after establishment is attributable to the conversion from a ploughed arable field to permanent grassland.
“An increase in SOM was expected because the experiment had been established on a ploughed arable field turned into permanent grassland.”
The paper relies on this without support.
The 13CO2 labelling experiment in the Ecotron facility is representative of root exudation effects on rhizospheric microorganisms in The Jena Experiment field plots.
“as indicated by a complementary 13 CO2 labelling experiment in the Ecotron facility in Montpellier (Methods).”
The paper relies on this without support.
The path model's causal structure is correctly specified and no unmeasured confounding affects the inferred relationships among plant diversity, root inputs, microbial activity and soil carbon storage.
“Path analyses revealed that plant species richness had direct positive effects on root carbon inputs as well as metabolic activity of soil microorganisms, measured as basal respiration (Fig. 3a,b; Methods).”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The lower carbon sequestration rates of herbaceous than woody crops after 11 years is a consequence of the effect of different factors such as carbon input dynamics from litter and roots.
“The lower C sequestration rates of herbaceous than woody crops after 11 years is a consequence of the effect of different factors such as C input dynamics from litter and roots, as explained above (Lange et al., 2015; Poeplau & Don, 2014; Zatta et al., 2014; Zimmermann et al., 2013).”
From Belowground biomass C outweighs soil organic C of perennial energy crops: Insights from a long‐term multispecies trial (2021) · cites this paper as Lange et al., 2015
The Results (p. 2) identify fine-root carbon inputs and microbial activity as drivers of soil carbon storage, and the Discussion (p. 4) describes turnover of root litter and exudates. But this was a nine-year grassland diversity experiment (p. 2); it does not establish that herbaceous crops sequester less carbon than woody crops after 11 years or explain that comparison through litter and root inputs. (LLM, conf=0.87)
Experimental — accuracy not yet measuredIn upper soil layers, carbon input from fine roots can trigger fast metabolic processes by soil microorganisms, thus accelerating soil organic carbon turnover (the 'priming effect').
“On the contrary, in upper soil layers C input from fine roots can trigger fast metabolic processes by soil microorganisms, thus accelerating SOC turnover, namely the 'priming effect' (Kuzyakov, 2002; Kuzyakov et al., 2000; Lange et al., 2015; Zatta et al., 2014).”
From Belowground biomass C outweighs soil organic C of perennial energy crops: Insights from a long‐term multispecies trial (2021) · cites this paper as Lange et al., 2015
Advanced biofuels face challenges to secure year-round, sustainable feedstock supply with quality that meets conversion specifications and to keep costs throughout the year reasonable.
“Like all biomass value chains, advanced biofuels face challenges to secure year-round, sustainable feedstock supply [153][154][155]with quality that meets the conversion specifications and keep costs throughout the year reasonable [156].”
From Advanced biofuels to decarbonise European transport by 2030: markets, challenges, and policies that impact their successful market uptake (2021) · cites this paper as [156]
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Labelling duration is reported as both three and four weeks
MinorThe discussion says the microbial community was sampled after a three-week label application, while the Methods say the monoliths were labelled for four weeks.
“the soil microbial community was sampled immediately after a 3-week label application”
“Monoliths with high (16 plant species) and low plant diversity (4 plant species) were labelled with 13 CO2 for 4 weeks.”
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Self-citationFunctional diversity of leaf nitrogen concentrations drives grassland carbon fluxes
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Self-citation
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Plant diversity enhances productivity and soil carbon storage
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Restoring Abandoned Farmland to Mitigate Climate Change on a Full Earth
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Do diverse cover crop mixtures perform better than monocultures? A systematic review
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- Authors
- Lange, M., Eisenhauer, N., Sierra, C. A., Bessler, H., Engels, C., Griffiths, R. I., Mellado-Vázquez, P. G., Malik, A. A., Roy, J., Scheu, S., Steinbeiss, S., Thomson, B. C., Trumbore, S. E., & Gleixner, G.
- Affiliations
- Max Planck Institute for Biogeochemistry (DE)
- German Centre for Integrative Biodiversity Research (DE)
- Humboldt-Universität zu Berlin (DE)
- UK Centre for Ecology & Hydrology (GB)
- Centre National de la Recherche Scientifique (FR)
- University of Göttingen (DE)
- Corresponding author
- Lange, Markus
- Gleixner, Gerd
- ORCID
- Venue
- Nature Communications
- Published
- 2015-04-07
- Volume
- 6
- Issue
- 1
- Pages
- 6707-6707
- ISSN
- 2041-1723 (ISSN-L)
- OpenAlex
- W2022423925
- Cited by
- 1,742
- References
- 69
- Citations by year
- 2026: 176 · 2025: 261 · 2024: 276 · 2023: 199 · 2022: 190 · 2021: 168 · 2020: 147 · 2019: 121 · 2018: 87 · 2017: 64 · 2016: 42 · 2015: 10
- FWCI
- 95.87
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Nature Communications (publishedVersion)
- PubMed (publishedVersion)
- NERC Open Research Archive (Natural Environment Research Council) (acceptedVersion)
- HAL (submittedVersion)
- RePEc: Research Papers in Economics (submittedVersion)
- eScholarship (California Digital Library) (submittedVersion, cc-by)
- GRO (submittedVersion, cc-by)
- MPG.PuRe (Max Planck Society) (submittedVersion)
- Funders
- Friedrich-Schiller-Universität Jena
- International Max Planck Research School for Environmental, Cellular and Molecular Microbiology
- Deutsche Forschungsgemeinschaft (FOR 456, FZT118)
- Agence Nationale de la Recherche (ANR-11-INBS-0001, ANR-11-INBS)
- Centre National de la Recherche Scientifique (ANR-11-INBS-0001)
- Deutsches Zentrum für integrative Biodiversitätsforschung Halle-Jena-Leipzig (FZT 118)
- Natural Environment Research Council (ceh020008, ceh020009)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Soil Science › Soil Carbon and Nitrogen Dynamics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Mycorrhizal Fungi and Plant Interactions
- Physical Sciences › Environmental Science › Nature and Landscape Conservation › Ecology and Vegetation Dynamics Studies
- Keywords
- plant diversity, soil carbon storage, soil microbial activity, soil microbial communities, carbon decomposition, grassland biodiversity, Jena Experiment, terrestrial carbon
- MeSH terms
- Carbon, Carbon Radioisotopes, Germany, Plants, Soil, Soil Microbiology, Ecosystem, Biodiversity, Grassland
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-10.
Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-10.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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