Biogeochemical Responses of Soil Organic Carbon and Enzymatic Activity to Elevation and Depth: Comparing Fixed Depth and Equivalent Soil Mass Approaches

Document Type : Research Paper

Authors

Department of Soil Science, Faculty of Agriculture, University of Zanjan, Zanjan, Iran

10.22059/ijswr.2025.399910.669990

Abstract

Soil organic carbon stocks (SOCs) and enzyme activities are key indicators for assessing ecosystem dynamics in mountainous regions. This study examined variations in SOCs and the activity of major enzymes along an altitudinal gradient (0–2400 m) and a depth gradient (0–100 cm) in mountain forests. Two approaches, fixed depth (FD) and equivalent soil mass (ESM), were applied to estimate SOCs. The results showed that SOCs were significantly influenced by altitude, depth, and their interaction (P ≤ 0.001). SOCs increased with altitude but decreased with depth; the highest and lowest values up to one meter depth were recorded at altitudes of 1800–2400 m and 0–600 m, respectively, representing a 1.14-fold increase in SOCs at higher elevations. Moreover, mean SOCs in deep layers (80–100 cm) showed about a 70% reduction compared with the surface layer (0–20 cm). The ESM method provided greater accuracy than the fixed-depth method (CV = 2.61% vs. 3.65%). The activities of β-glucosidase, urease, and arylsulfatase increased with altitude but decreased with depth, whereas cellulase, acid phosphatase, and alkaline phosphatase decreased with both factors. In contrast, dehydrogenase (DHA) activity increased with depth and exhibited a negative correlation with SOCs (r = –0.92). Additionally, SOCs were strongly correlated with microbial biomass carbon (r = 0.99) and enzyme activities (r ≥ 0.82), while its depth distribution remained consistent across the entire altitudinal gradient. These findings underscore the importance of jointly considering soil depth and altitude in SOC modeling and the sustainable management of mountain ecosystems.

Keywords

Main Subjects


Introduction

Soil Organic Carbon (SOC) is a critical component of terrestrial carbon pools and plays a key role in soil health, productivity, and climate change mitigation. In mountainous forest ecosystems, SOC stocks and related microbial processes are shaped by topographic variations, particularly elevation and soil depth. These factors jointly influence microclimatic conditions, vegetation dynamics, and organic matter decomposition. Enzyme-mediated biogeochemical processes serve as sensitive indicators of soil microbial activity and are tightly linked to carbon cycling. However, the combined effects of elevation and depth on SOC and enzyme activity have not been comprehensively studied, especially using both fixed-depth and equivalent soil mass (ESM) approaches. This study aimed to evaluate SOC stocks and key enzyme activities along an elevation gradient and across soil profiles in a mountainous forest ecosystem.

Methods

The study was conducted in a mountainous forest region covering an elevation range from 0 to 2400 meters above sea level. Soil samples were collected along this gradient using two methods: the conventional fixed-depth method and the Equivalent Soil Mass (ESM) approach, which accounts for differences in soil bulk density to provide more accurate carbon stock estimations. Samples were taken from five soil depth intervals: 0–20 cm, 20–40 cm, 40–60 cm, 60–80 cm, and 80–100 cm. SOC was measured using standard dry combustion methods. Microbial activity was assessed by analyzing the activity of key soil enzymes: β-glucosidase, urease, arylsulfatase, cellulase, acid phosphatase, alkaline phosphatase, and dehydrogenase. Statistical analyses including ANOVA and Pearson correlation were applied to evaluate the effects of elevation and soil depth on SOC stocks and enzyme activities.

Results

SOC stocks were significantly affected by elevation, soil depth, and their interaction (P ≤ 0.001). Soil depth had the strongest influence, with SOC stocks decreasing by approximately 70% from the surface layer (0–20 cm) to the deepest layer (80–100 cm). In contrast, SOC stocks increased by 1.14 times from the lowest to the highest elevation. The ESM method provided higher sensitivity and accuracy than the fixed-depth method, with a lower coefficient of variation (CV = 2.61% vs. 3.65%). It was more effective in detecting significant SOC differences across elevation and precipitation gradients. Enzyme activities responded differently to changes in elevation and depth. Activities of β-glucosidase, urease, and arylsulfatase increased significantly with elevation but declined with increasing soil depth. Conversely, cellulase, acid phosphatase, and alkaline phosphatase activities showed a decreasing trend along both elevation and depth gradients. Dehydrogenase activity exhibited an opposite pattern, increasing up to 49 times with depth and showing a strong negative correlation with SOC (R² = 0.83), suggesting microbial adaptation to carbon-limited conditions in deeper layers. SOC also showed a strong positive correlation with microbial biomass carbon (r = 0.99) and enzyme activity (r ≥ 0.82). Despite the increase in total SOC stocks with elevation, the relative distribution of carbon with soil depth remained consistent. This suggests structural stability in vegetation cover and root distribution across the elevation gradient.

Conclusions

This study highlights the importance of accounting for both elevation and soil depth when assessing SOC dynamics in mountainous forest ecosystems. The Equivalent Soil Mass (ESM) method proved to be a more accurate and sensitive tool for estimating SOC stocks under varying topographic and climatic conditions. The contrasting enzyme activity patterns underscore the complexity of microbial responses to environmental gradients. While SOC stocks increased with elevation, the vertical distribution of carbon remained stable, likely due to consistent vegetation structure. These findings emphasize the need to integrate both vertical and elevational dimensions in carbon modeling and land management strategies, particularly in topographically complex and climate-sensitive regions.

Author Contributions

Elham Jozedaemi: Writing – original draft, Methodology, Formal analysis, Investigation, Resources.

Ahmad Golchin: Writing – review and editing, Supervision, Methodology, Conceptualization, Resources.

Mehran Misaghi: Writing – review and editing, Investigation, Resources, Software, Formal analysis.

Data Availability Statement

Data available on request from the authors.

 

Acknowledgements

The authors express their gratitude to the University of Zanjan, Zanjan, Iran, for its financial and technical support, which facilitated the completion of this study.

Conflict of interest

The author declares no conflict of interest.

Afreen, T., Singh, H., & Singh, J. S. (2019). Influence of changing patterns of precipitation and temperature on tropical soil ecosystem. Tropical ecosystems: Structure, functions and challenges in the face of global change, 11-26. https://doi.org/10.1007/978-981-13-8249-9_2
Anderson, J. P., & Domsch, K. H. (1978). A physiological method for the quantitative measurement of microbial biomass in soils. Soil biology and biochemistry10 (3), 215-221. https://doi.org/10.1016/0038-0717(78)90099-8
Barry, R. G. (2008). Mountain weather and climate (pp. xxiv+-506). https://www.cabidigitallibrary.org/doi/full/10.5555/20093032476
Batjes, N. H. (1996). Total carbon and nitrogen in the soils of the world. European journal of soil science47(2), 151-163. https://doi.org/10.1111/j.1365-2389.1996.tb01386.x
Batjes, N. H. (2014). Total carbon and nitrogen in the soils of the world. European journal of soil science65 (1), 10-21. https://doi.org/10.1111/ejss.12114_2
Bell, C. W., Fricks, B. E., Rocca, J. D., Steinweg, J. M., McMahon, S. K., & Wallenstein, M. D. (2013). High-throughput fluorometric measurement of potential soil extracellular enzyme activities. Journal of visualized experiments: JoVE, (81), 50961. https://doi.org/10.3791/50961
Blake, G. R. (1965). Bulk density. Methods of soil analysis: Part 1 physical and mineralogical properties, including statistics of measurement and sampling9, 374-390. https://doi.org/10.2134/agronmonogr9.1.c30
Canadell, J., Jackson, R. B., Ehleringer, J. B., Mooney, H. A., Sala, O. E., & Schulze, E. D. (1996). Maximum rooting depth of vegetation types at the global scale. Oecologia108, 583-595. https://doi.org/10.1007/BF00329030
Conant, R. T., Ryan, M. G., Ågren, G. I., Birge, H. E., Davidson, E. A., Eliasson, P. E., ... & Bradford, M. A. (2011). Temperature and soil organic matter decomposition rates–synthesis of current knowledge and a way forward. Global change biology17 (11), 3392-3404. https://doi.org/10.1111/j.1365-2486.2011.02496.x
Cotrufo, M. F., Wallenstein, M. D., Boot, C. M., Denef, K., & Paul, E. (2013). The M icrobial E fficiency‐M atrix S tabilization (MEMS) framework integrates plant litter decomposition with soil organic matter stabilization: do labile plant inputs form stable soil organic matter?. Global change biology19 (4), 988-995. https://doi.org/10.1111/gcb.12113
Crowther, T. W., Todd-Brown, K. E., Rowe, C. W., Wieder, W. R., Carey, J. C., Machmuller, M. B., ... & Bradford, M. A. (2016). Quantifying global soil carbon losses in response to warming. Nature540 (7631), 104-108. https://doi.org/10.1038/nature20150
Djukic, I., Zehetner, F., Tatzber, M., & Gerzabek, M. H. (2010). Soil organic‐matter stocks and characteristics along an Alpine elevation gradient. Journal of Plant Nutrition and Soil Science173 (1), 30-38. https://doi.org/10.1002/jpln.200900027
Don, A., Rödenbeck, C., & Gleixner, G. (2013). Unexpected control of soil carbon turnover by soil carbon concentration. Environmental Chemistry Letters11, 407-413. https://doi.org/10.1007/s10311-013-0433-3
Egamberdieva, D., Renella, G., Wirth, S., & Islam, R. (2010). Enzyme activities in the rhizosphere of plants. In Soil enzymology (pp. 149-166). Berlin, Heidelberg: Springer Berlin Heidelberg. https://doi.org/10.1007/978-3-642-14225-3_8
Fierer, N., & Jackson, R. B. (2006). The diversity and biogeography of soil bacterial communities. Proceedings of the National Academy of Sciences103 (3), 626-631. https://doi.org/10.1073/pnas.0507535103
Garcia, C., Hernandez, T., & Costa, F. (1997). Potential use of dehydrogenase activity as an index of microbial activity in degraded soils. Communications in soil science and plant analysis28 (1-2), 123-134. https://doi.org/10.1080/00103629709369777
Garcia-Pausas, J., Casals, P., Camarero, L., Huguet, C., Sebastia, M. T., Thompson, R., & Romanya, J. (2007). Soil organic carbon storage in mountain grasslands of the Pyrenees: effects of climate and topography. Biogeochemistry82, 279-289. https://doi.org/10.1007/s10533-007-9071-9
García‐Palacios, P., Vandegehuchte, M. L., Shaw, E. A., Dam, M., Post, K. H., Ramirez, K. S., ... & Wall, D. H. (2015). Are there links between responses of soil microbes and ecosystem functioning to elevated CO 2, N deposition and warming? A global perspective. Global Change Biology21 (4), 1590-1600. https://doi.org/10.1111/gcb.12788
Gholami, R. (2019). Mineralogical evolution of Inceptisols and Alfisols at two elevation levels in Arasbaran forests [Master’s thesis, University of Tabriz].
Goidts, E., Van Wesemael, B., & Crucifix, M. (2009). Magnitude and sources of uncertainties in soil organic carbon (SOC) stock assessments at various scales. European Journal of Soil Science60 (5), 723-739. https://doi.org/10.1111/j.1365-2389.2009.01157.x
Guo, M., Zhao, B., Wen, Y., Hu, J., Dou, A., Zhang, Z., ... & Zhu, J. (2022). Elevational pattern of soil organic carbon release in a Tibetan alpine grassland: Consequence of quality but not quantity of initial soil organic carbon. Geoderma428, 116148. https://doi.org/10.1016/j.geoderma.2022.116148
Guo, L. B., & Gifford, R. M. (2002). Soil carbon stocks and land use change: a meta analysis. Global change biology8 (4), 345-360. https://doi.org/10.1046/j.1354-1013.2002.00486.x
Gutiérrez-Girón, A., Díaz-Pinés, E., Rubio, A., & Gavilán, R. G. (2015). Both altitude and vegetation affect temperature sensitivity of soil organic matter decomposition in Mediterranean high mountain soils. Geoderma237, 1-8. https://doi.org/10.1016/j.geoderma.2014.08.005
Hagedorn, F., Moeri, A., Walthert, L., & Zimmermann, S. (2010). Kohlenstoff in Schweizer Waldböden–bei Klimaerwärmung eine potenzielle CO2-Quelle| Soil organic carbon in Swiss forest soils–a potential CO2 source in a warming climate. Schweizerische Zeitschrift für Forstwesen161 (12), 530-535. https://doi.org/10.3188/szf.2010.0530
Hofmann, E. (1965). Urease. In Methods of enzymatic analysis (pp. 913-916). Academic Press. https://doi.org/10.1016/B978-0-12-395630-9.50163-8
Huluka, G., & Miller, R. (2014). Particle size determination by hydrometer method. Southern Cooperative Series Bulletin419, 180-184. https://aesl.ces.uga.edu/sera6/PUB/MethodsManualFinalSERA6.pdf#page=188
Ipcc, I. P. C. C. (2006). Guidelines for national greenhouse gas inventories. Prepared by the National Greenhouse Gas Inventories Programme. Eggleston HS, Buendia L, Miwa K, Ngara T, Tanabe K, editors. Published: IGES, Japan. https://www.ipcc-nggip.iges.or.jp/meeting/pdfiles/Washington_Report.pdf
Jackson, R. B., Lajtha, K., Crow, S. E., Hugelius, G., Kramer, M. G., & Piñeiro, G. (2017). The ecology of soil carbon: pools, vulnerabilities, and biotic and abiotic controls. Annual review of ecology, evolution, and systematics48 (1), 419-445. https://doi.org/10.1146/annurev-ecolsys-112414-054234
Kallenbach, C. M., Grandy, A. S., Frey, S. D., & Diefendorf, A. F. (2015). Microbial physiology and necromass regulate agricultural soil carbon accumulation. Soil Biology and Biochemistry91, 279-290. https://doi.org/10.1016/j.soilbio.2015.09.005
Kleber, M., Sollins, P., & Sutton, R. (2007). A conceptual model of organo-mineral interactions in soils: self-assembly of organic molecular fragments into zonal structures on mineral surfaces. Biogeochemistry85, 9-24. https://doi.org/10.1007/s10533-007-9103-5
Koch, O., Tscherko, D., & Kandeler, E. (2007). Temperature sensitivity of microbial respiration, nitrogen mineralization, and potential soil enzyme activities in organic alpine soils. Global Biogeochemical Cycles21 (4). https://doi.org/10.1029/2007GB002983
Kong, J., He, Z., Chen, L., Zhang, S., Yang, R., & Du, J. (2022). Elevational variability in and controls on the temperature sensitivity of soil organic matter decomposition in alpine forests. Ecosphere13 (4), e4010. https://doi.org/10.1002/ecs2.4010
Kögel‐Knabner, I., Guggenberger, G., Kleber, M., Kandeler, E., Kalbitz, K., Scheu, S., ... & Leinweber, P. (2008). Organo‐mineral associations in temperate soils: Integrating biology, mineralogy, and organic matter chemistry. Journal of Plant Nutrition and Soil Science171 (1), 61-82. https://doi.org/10.1002/jpln.200700048
Körner, C. (2007). The use of ‘altitude’in ecological research. Trends in ecology & evolution22 (11), 569-574. https://doi.org/10.1016/j.tree.2007.09.006
Lehmann, J., & Kleber, M. (2015). The contentious nature of soil organic matter. Nature528  (7580), 60-68. https://doi.org/10.1038/nature16069
Leifeld, J., & Fuhrer, J. (2005). The temperature response of CO 2 production from bulk soils and soil fractions is related to soil organic matter quality. Biogeochemistry75, 433-453. https://doi.org/10.1007/s10533-005-2237-4
Li, X., Xie, J., Zhang, Q., Lyu, M., Xiong, X., Liu, X., ... & Yang, Y. (2020). Substrate availability and soil microbes drive temperature sensitivity of soil organic carbon mineralization to warming along an elevation gradient in subtropical Asia. Geoderma364, 114198. https://doi.org/10.1016/j.geoderma.2020.114198
Lloyd, J., & Taylor, J. A. (1994). On the temperature dependence of soil respiration. Functional ecology, 315-323. https://doi.org/10.2307/2389824
Luo, Z., Feng, W., Luo, Y., Baldock, J., & Wang, E. (2017). Soil organic carbon dynamics jointly controlled by climate, carbon inputs, soil properties and soil carbon fractions. Global change biology23 (10), 4430-4439. https://doi.org/10.1111/gcb.13767
Mathieu, J. A., Hatté, C., Balesdent, J., & Parent, É. (2015). Deep soil carbon dynamics are driven more by soil type than by climate: a worldwide meta‐analysis of radiocarbon profiles. Global change biology21 (11), 4278-4292. https://doi.org/10.1111/gcb.13012
Margalef, O., Sardans, J., Fernández-Martínez, M., Molowny-Horas, R., Janssens, I. A., Ciais, P., ... & Penuelas, J. (2017). Global patterns of phosphatase activity in natural soils. Scientific reports7 (1), 1337. https://doi.org/10.1038/s41598-017-01418-8
Martín, M. Á., Reyes, M., & Taguas, F. J. (2017). Estimating soil bulk density with information metrics of soil texture. Geoderma287, 66-70. https://doi.org/10.1016/j.geoderma.2016.09.008
Merino, C., Godoy, R., & Matus, F. (2016). Soil enzymes and biological activity at different levels of organic matter stability. Journal of soil science and plant nutrition16 (1), 14-30. http://dx.doi.org/10.4067/S0718-95162016005000002
Mishra, U., Ussiri, D. A., & Lal, R. (2010). Tillage effects on soil organic carbon storage and dynamics in Corn Belt of Ohio USA. Soil and Tillage Research107 (2), 88-96. https://doi.org/10.1016/j.still.2010.02.005
Montgomery, D. R. (2007). Soil erosion and agricultural sustainability. Proceedings of the National Academy of Sciences104 (33), 13268-13272. https://doi.org/10.1073/pnas.0611508104
Nogués-Bravo, D., Araújo, M. B., Errea, M. P., & Martínez-Rica, J. P. (2007). Exposure of global mountain systems to climate warming during the 21st Century. Global environmental change17 (3-4), 420-428. https://doi.org/10.1016/j.gloenvcha.2006.11.007
Öhlinger, H., & Von Mersi, W. (1996). Enzymes involved in intracellular metabolism. In Methods in soil biology (pp. 235-245). Berlin, Heidelberg: Springer Berlin Heidelberg. https://doi.org/10.1007/978-3-642-60966-4_15
Pan, Y., Yu, S. S., Xiao, Z. C., Min, Y., Tian, T., Zheng, Y. M., ... & Yu, H. Q. (2023). Re-evaluation and modification of dehydrogenase activity tests in assessing microbial activity for wastewater treatment plant operation. Water Research246, 120737. https://doi.org/10.1016/j.watres.2023.120737
Paustian, K., Lehmann, J., Ogle, S., Reay, D., Robertson, G. P., & Smith, P. (2016). Climate-smart soils. Nature532 (7597), 49-57. https://doi.org/10.1038/nature17174
Powlson, D. S., Whitmore, A. P., & Goulding, K. W. (2011). Soil carbon sequestration to mitigate climate change: a critical re‐examination to identify the true and the false. European journal of soil science62 (1), 42-55. https://doi.org/10.1111/j.1365-2389.2010.01342.x
Prescott, C. E. (2010). Litter decomposition: what controls it and how can we alter it to sequester more carbon in forest soils?. Biogeochemistry101, 133-149. https://doi.org/10.1007/s10533-010-9439-0
Raffeld, A. M., Bradford, M. A., Jackson, R. D., Rath, D., Sanford, G. R., Tautges, N., & Oldfield, E. E. (2024). The importance of accounting method and sampling depth to estimate changes in soil carbon stocks. Carbon Balance and Management19 (1), 2. https://doi.org/10.1186/s13021-024-00249-1
Rezaei, H., Jsfarzadeh, A. A., Alijanpour, A., Shahbazi, F., & Valizadeh Kamran, K. (2017). Genetically evolution of Arasbaran forests soils along altitudinal transects of Kaleybar Chai Sofla Sub-Basin. Water and Soil Science26 (4.1), 151-166. https://water-soil.tabrizu.ac.ir/article_5864.html?lang=en
Rezaei, H., Jafarzadeh, A. A., Alijanpour, A., Shahbazi, F., & Valizadeh Kamran, K. (2020). Soil Organic Matter Condition in Forest Stands of Arasbaran. Water and Soil34 (1), 115-127. https://doi.org/10.22067/jsw.v34i1.80633
Rumpel, C., & Kögel-Knabner, I. (2011). Deep soil organic matter—a key but poorly understood component of terrestrial C cycle. Plant and soil338, 143-158. https://doi.org/10.1007/s11104-010-0391-5
Rustad, L. E. J. L., Campbell, J., Marion, G., Norby, R., Mitchell, M., Hartley, A., ... & Gcte-News. (2001). A meta-analysis of the response of soil respiration, net nitrogen mineralization, and aboveground plant growth to experimental ecosystem warming. Oecologia126, 543-562. https://doi.org/10.1007/s004420000544
Schimel, J. P., & Schaeffer, S. M. (2012). Microbial control over carbon cycling in soil. Frontiers in microbiology3, 348. https://doi.org/10.3389/fmicb.2012.00348
Schinner, F., Öhlinger, R., Kandeler, E., & Margesin, R. (1996). Enzymes involved in carbon metabolism. In Methods in Soil Biology (pp. 185-207). Berlin, Heidelberg: Springer Berlin Heidelberg. https://doi.org/10.1007/978-3-642-60966-4_12
Schmidt, M. W., Torn, M. S., Abiven, S., Dittmar, T., Guggenberger, G., Janssens, I. A., ... & Trumbore, S. E. (2011). Persistence of soil organic matter as an ecosystem property. Nature478 (7367), 49-56. https://doi.org/10.1038/nature10386
Schuur, E. A., Bockheim, J., Canadell, J. G., Euskirchen, E., Field, C. B., Goryachkin, S. V., ... & Zimov, S. A. (2008). Vulnerability of permafrost carbon to climate change: Implications for the global carbon cycle. BioScience58 (8), 701-714. https://doi.org/10.1641/B580807
Sinsabaugh, R. L., Lauber, C. L., Weintraub, M. N., Ahmed, B., Allison, S. D., Crenshaw, C., ... & Zeglin, L. H. (2008). Stoichiometry of soil enzyme activity at global scale. Ecology letters11 (11), 1252-1264. https://doi.org/10.1111/j.1461-0248.2008.01245.x
Sinsabaugh, R. L., Hill, B. H., & Follstad Shah, J. J. (2009). Ecoenzymatic stoichiometry of microbial organic nutrient acquisition in soil and sediment. Nature462 (7274), 795-798. https://doi.org/10.1038/nature08632
Smith, P. (2008). Land use change and soil organic carbon dynamics. Nutrient Cycling in Agroecosystems81, 169-178. https://doi.org/10.1007/s10705-007-9138-y
Souza, C. R., Mariano, R. F., Maia, V. A., Pompeu, P. V., Dos Santos, R. M., & Fontes, M. A. L. (2023). Carbon stock and uptake in the high-elevation tropical montane forests of the threatened Atlantic Forest hotspot: Ecosystem function and effects of elevation variation. Science of The Total Environment882, 163503. https://doi.org/10.1016/j.scitotenv.2023.163503
Steinweg, J. M., Dukes, J. S., Paul, E. A., & Wallenstein, M. D. (2013). Microbial responses to multi-factor climate change: effects on soil enzymes. Frontiers in microbiology4, 146. https://doi.org/10.3389/fmicb.2013.00146
Stockmann, U., Adams, M. A., Crawford, J. W., Field, D. J., Henakaarchchi, N., Jenkins, M., ... & Zimmermann, M. (2013). The knowns, known unknowns and unknowns of sequestration of soil organic carbon. Agriculture, Ecosystems & Environment164, 80-99. https://doi.org/10.1016/j.agee.2012.10.001
Stone, M. M., DeForest, J. L., & Plante, A. F. (2014). Changes in extracellular enzyme activity and microbial community structure with soil depth at the Luquillo Critical Zone Observatory. Soil Biology and Biochemistry75, 237-247. https://doi.org/10.1016/j.soilbio.2014.04.017
Sundqvist, M. K., Sanders, N. J., & Wardle, D. A. (2013). Community and ecosystem responses to elevational gradients: processes, mechanisms, and insights for global change. Annual review of ecology, evolution, and systematics44 (1), 261-280. https://doi.org/10.1146/annurev-ecolsys-110512-135750
Tashi, S., Singh, B., Keitel, C., & Adams, M. (2016). Soil carbon and nitrogen stocks in forests along an altitudinal gradient in the eastern Himalayas and a meta‐analysis of global data. Global change biology22 (6), 2255-2268. https://doi.org/10.1111/gcb.13234
Tunlid, A., & White, D. C. (2021). Biochemical analysis of biomass, community structure, nutritional status, and metabolic activity of microbial communities in soil. In Soil biochemistry (pp. 229-262). CRC Press. https://doi.org/10.1201/9781003210207
VandenBygaart, A. J., & Angers, D. A. (2006). Towards accurate measurements of soil organic carbon stock change in agroecosystems. Canadian Journal of Soil Science86 (3), 465-471. https://doi.org/10.4141/S05-106
Von Haden, A. C., Yang, W. H., & DeLucia, E. H. (2020). Soils' dirty little secret: Depth‐based comparisons can be inadequate for quantifying changes in soil organic carbon and other mineral soil properties. Global Change Biology26 (7), 3759-3770. https://doi.org/10.1111/gcb.15124
Walkley, A., & Black, I. A. (1934). An examination of the Degtjareff method for determining soil organic matter, and a proposed modification of the chromic acid titration method. Soil science37 (1), 29-38. https://journals.lww.com/soilsci/toc/1934/01000
Wendt, J. W., & Hauser, S. (2013). An equivalent soil mass procedure for monitoring soil organic carbon in multiple soil layers. European Journal of Soil Science64 (1), 58-65. https://doi.org/10.1111/ejss.12002
Wiesmeier, M., Burmeister, J., Garcia-Franco, N., & Sümmerer, M. (2024). Soil Organic Carbon and Nitrogen Changes in Agricultural Soils of Bavaria Between 1986 and 2016 as Driven by Management and Climate Change. Available at SSRN 4901937. https://dx.doi.org/10.2139/ssrn.4901937
Yoo, K., Amundson, R., Heimsath, A. M., & Dietrich, W. E. (2006). Spatial patterns of soil organic carbon on hillslopes: Integrating geomorphic processes and the biological C cycle. Geoderma130 (1-2), 47-65. https://doi.org/10.1016/j.geoderma.2005.01.008
Zhang, R., & Wienhold, B. J. (2002). The effect of soil moisture on mineral nitrogen, soil electrical conductivity, and pH. Nutrient cycling in Agroecosystems63, 251-254. https://doi.org/10.1023/A:1021115227884