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Weight and dimensions of fruit and stoneThe mean weight of a fruit from a particular tree ranged from 21.25 ± 4.26 to 58.26 ± 10.44 g, with the weight of the heaviest mean fruit weight being 2.74 times that of the lightest. Similarly, the mean stone weight ranged from 8.99 ± 2.35 to 20.32 ± 3.14 g, with a 2.26 times difference between the heaviest and lightest stones (Table 2). There were significant differences (p  More

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Bed bugsFour bed bug populations (one laboratory strain and three collected from infested homes) were used in this study (Table 1). All populations were reared in the laboratory as described by DeVries et al.28. Briefly, bed bugs were maintained in 168 cm3 plastic containers on paper substrate at 25 °C, 50% relative humidity, and a photoperiod of 12 h:12 h (Light:Dark). Bed bugs were fed defibrinated rabbit blood (Hemostat Laboratories, Dixon, CA, USA) weekly using an artificial feeding system. This system maintained blood at 35 °C by circulating water through custom-made water-jacketed glass feeders. An artificial membrane (plant budding tape, A.M. Leonared, Piqua, OH, USA) was stretched over the bottom of each glass feeder, containing the blood while simultaneously allowing bed bugs to feed through it. In all experiments, adult males starved for 7–10 days were used. All populations were used for documenting responses to human skin swabs. The WS population was used for bioassays with various human volunteers and hexane extracted swabs, and the JC population was used for testing various lipids.Table 1 Bed bug populations used in this study.Full size tableSkin swab collectionThe North Carolina State University Institutional Review Board approved this study (IRB #14173). Informed consent was obtained from all human participants, and all the methods were performed according to the relevant guidelines and regulations. Six human volunteers (3 males, 3 females) ranging from 25 to 50 years old representing several ethnicities (white/Caucasian, Hispanic, Asian) provided samples for this project. Skin swabs were collected following the exact methods outlined by DeVries et al.16. In our 2019 study, these swabs were reported to attract bed bugs independent of other cues in Y-tube olfactometer assays. Briefly, participants were asked to follow a standard operating procedure, which was reviewed with them prior to sample collection. Before collecting skin swabs, participants were asked to not to eat ‘spicy’ food for at least 24 h, take a morning shower, avoid the use of deodorant and cosmetics after showering, and avoid strenuous physical activity. Skin swabs were collected 4–8 h after showering. Hands were washed with water only before lifting filter paper. Swabs were collected using 4.5 cm diameter filter paper discs (#1; Whatman plc, Madistone, United Kingdom). Both sides of a single filter paper disc were rubbed over the left arm from hand to armpit for 12 s, left leg from lower thigh to ankle for 12 s, and left armpit for 6 s. This procedure was repeated on the right side using a new filter paper disc, so that two samples were collected during each swabbing session. The skin swab samples were then stored in glass vials at − 20 °C, and used within one month of collection. The swabs from all human volunteers were used to compare participants and establish that bed bugs responded similarly to all, and participant A’s skin swabs were used for all subsequent bioassays.Two-choice arrestment bioassaysTwo-choice bioassays were conducted in plastic Petri dishes of 6 cm diameter (Corning Life Sciences, Durham, NC, USA) (Fig. 1). The bottom surface of each Petri dish was roughened so that bed bugs could freely move about the arena. Two tents (3 × 1.5 cm) were created using filter paper (Whatman #1). One tent served as the control tent, and the other served as the treatment tent. Control tents were either untreated (nothing added) or treated with hexane only. Treatment tents were either made directly from human odor swabs, treated with human odor extract (in hexane), or treated with a specific compound (in hexane). Tents were allowed 60 min to acclimate to room conditions and allow for the solvent to evaporate prior to initiating bioassays. The positions of tents (treatment and control) were alternated to account for any side-bias.Figure 1Two-choice behavioral assay (top-view) consisting of two equal size paper shelter tents. A clean filter paper (control) was always paired with a treated filter paper that either represented a human skin swab, hexane extract of swabbed paper, SPE fraction of human skin swab extract, or authentic TAGs. A single male bed bug was introduced into the center of each arena and allowed to select a tent to arrest under.Full size imageAdult male bed bugs were housed in individual vials for 24 h prior to each experiment. A single adult male bed bug was released in the middle of the arena 5 h into the scotophase, by transferring it on its harborage. The harborage material was removed immediately after the bed bug moved off of it (the harborage). Bed bugs were allowed the remaining 7 h of the scotophase to freely move around the arena, with their final position reported 3 h into the photophase. Bed bugs that were in contact with the filter paper with any part of their body were recorded as making a choice (i.e. arrestment state); others not in contact with either filter paper tent were recorded as non-responders, reported in the figures, but not used in data analysis. It should be noted that momentary pauses in movement (feeding or other behaviors) are not referred to as arrestment in this study. In total, 15–39 replicates were performed for each experiment (reported for each bioassay).Bioassays with human skin swabsBioassays with human skin swabs were performed to understand if bed bug arrestment behavior (1) differed among different bed bug populations, and (2) influenced by different host odors. Skin swabs were removed from the freezer, equilibrated to room temperature, divided into three equal parts and trimmed to a rectangular shape corresponding to the size of a shelter tent (Fig. 1). Skin swabs from participant A were used to evaluate the responses of four bed bug populations (Table 1). Skin swabs from all participants A–F were used to evaluate the robustness of our findings across multiple human hosts.Skin swab extraction and fractionationSkin swabs collected from volunteer A were pooled and extracted in hexane. Extraction procedures were carried out sequentially by placing a single skin swab into a 20 ml glass vial containing 5 ml of hexane, vortexing for 30 s, then moving the filter paper to a new 20 ml vial containing 5 ml of hexane and repeating the process. Three sequential extractions were performed for each skin swab, and a minimum of 10 skin swabs (collected over several days) were used for each extraction. After all skin swabs were extracted, all sequential hexane extracts were combined and concentrated to a final concentration of one skin swab equivalent per 300 µl, or one bioassay equivalent (BE) per 100 µl (since each swab was used for 3 bioassays; see “Bioassays with human skin swabs” for more information on the size used for each bioassay). Control swabs were also extracted. These swabs were treated identically to the skin swabs, except they did not contact human skin.To determine what compound classes were responsible for the observed behavior, hexane extracts were fractionated using solid phase extraction (SPE). Extracted samples were concentrated to 1 BE/10 µl hexane, then loaded onto a 1 g silica SPE column (6 ml total volume; J.T. Baker, Phillipsburg, NJ, USA). The column was eluted with the following solvents (4 ml of each, each repeated twice sequentially): hexane, 2% ether (in hexane), 5% ether (in hexane), 10% ether (in hexane), 20% ether (in hexane), 50% ether (in hexane), 100% ether, ethyl acetate, and methanol (all solvents acquired from Sigma Aldrich, St. Louis, MO, USA). Each solvent fraction was then concentrated to a final concentration of 1 BE/100 µl and stored at − 20 °C.Bioassays with extracted and fractionated human skin swabsFor all extraction and fractionation bioassays, filter paper tents were cut to a size of 3 cm × 1.5 cm (Fig. 1) and treated with 100 µl (1 BE) of extracted or fractionated human skin swabs (50 µl on each side). A dose–response bioassay was run first to determine if the compounds responsible for bed bug arrestment responses could be extracted and at what concentration (BE) they were behaviorally active. Dilutions were made in hexane, with control tents receiving extracts of control filter paper. At least 20 replicates were conducted for each concentration. After validating an appropriate BE that could be used in future experiments, SPE fractions were diluted in hexane to 0.1 BE and applied to filter paper tents as previously described (50 µl per side). A minimum of 15 replicates were conducted for each fraction to identify behaviorally active fractions.Compound identificationTo better understand what classes of compounds were present in behaviorally active fractions, we conducted thin layer chromatography (TLC) with known standards. A flexible, silica (250 µm) TLC plate (Whatman) was placed into a glass chamber containing a solvent layer of 1.5 cm. The plate was cleaned twice with acetone, then standards (triglyceride [TAG], wax ester, squalene) and samples (fractions) were each loaded into separate lanes. The plate was developed twice in 10% ether (in hexane), then visualized non-destructively with iodine.In addition, behaviorally active fractions were further evaluated for their composition with GC–MS and LC–MS. GC–MS was employed to analyze free fatty acids, squalene, and cholesterol29, whereas LC–MS was employed to characterize the intact skin lipids as previously described30. Samples were analyzed with a GC 7890A coupled to the MS 5975 VL analyzer (Agilent Technologies, CA, USA) following derivatization. Briefly, 50 µL of the extract dissolved in isopropanol were dried under nitrogen and derivatized with 100 µL BSTFA containing 1% trimethylchlorosilane (TCMS) in pyridine to generate the trimethylsilyl (TMS) derivatives at 60 °C for 60 min. GC separation was performed with a 30 m × 0.250 mm (i.d.) × 0.25 µm film thickness DB-5MS fused silica column (Agilent). Helium was used as the carrier gas. Samples were acquired in scan mode by means of electron impact (EI) MS.Liquid-chromatography coupled to the MS analyzer by means of an electrospray interface (ESI) was used to determine abundance and ESI tandem MS of non-volatile lipids as previously described29,30. LC separation was performed with a reverse phase Zorbax SB-C8 column (2.1 × 100 mm, 1.8 μm particle size, Agilent). Data were acquired in the positive ion mode at unit mass resolving power by scanning ions between m/z 100 and 1000 with G6410A series triple quadrupole (QqQ) (Agilent). LC runs and MS spectra were processed with the Mass Hunter software (B.09.00 version).Bioassays with triglyceridesAfter determining that TAGs were prominent compounds in bioactive skin swab fractions, commercially available TAGs were evaluated for behavioral activity. Filter paper tents were treated with 100 µl of hexane (50 µl to each side) containing TAG standards. First, tripalmitin (16:0/16:0/16:0) (Sigma-Aldrich) was evaluated in a dose–response fashion (60 µg to 0.6 µg) to determine what level of TAG was appropriate for bioassays. The upper level of testing was set at 60 µg as a conservative estimate of the amount of TAGs bed bugs may be exposed to, based on calculations of our arena size and previous reports of TAGs on human skin and sebum. Specifically, previous reports documented that 1.5 mg of sebum could be passively collected using Sebutape from an area of 4.7 cm230,31. Because TAGs typically constitute 60% of human sebum32, it is reasonable to assume that passive collection of sebum can result in  > 190 µg/cm2 of TAGs in a short amount of time (30 min). Our sampling methods involved swabbing rather than passive collection, but our use of 60 µg over a 9 cm2 (two sides of 4.5 cm2) shelter tent (6.67 µg/cm2) is a low-estimate of the amount of TAGs collected (although this was not directly measured in the current study). Other TAGs that we tested at a concentration of 60 µg per 9 cm2 included the saturated TAGs trimyristin (14:0/14:0/14:0) and tristearin (18:0/18:0/18:0) and the unsaturated TAGs triolein (18:1/18:1/18:1), trilinolein (18:2/18:2/18:2), and trilinolenin (18:3/18:3/18:3) (all from Sigma-Aldrich). A minimum of 30 replicates were conducted with each TAG.Statistical analysisA Chi-square goodness of fit test was used to compare the responses of bed bugs to control versus treated tents in all two-choice bioassays, with the null hypothesis that if bed bugs do not respond differentially to treated tents they should display a 1:1 preference ratio for both sides of the assay. All tests were conducted in SPSS Version 26 (IBM Corp., Armonk, NY). More

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Site characteristics and experimental designIn this study, agricultural soils with five SOM contents were collected in 2015 from the following three different locations with the same climate type (the moderate temperate continental climate) in Northeast China (Table S3 and Fig. 1): Bei’an (BA), Hailun (HL), and Dehui (DH). Their MAT and MAP range from 1.0 to 4.4 and 520 to 550, respectively. After collection, the samples were transported to the Hailun Agricultural Ecological Experimental Station (HL), where the samples were packed into the same PVC tubes. Moving the soil from these three initial sampling points to the HL may have had some influence on the microbes, but compared with longer-distance soil translocation across different climatic zones, the HL site can be regarded as an in situ site that reflects the original climatic conditions. The SOM contents were 2%, 3%, 5%, 7%, and 9% (equivalent to 10, 18, 28, 36, and 56 g C kg−1 soil−1, respectively), and all the soils were classified as Mollisols according to the FAO classification. Here, we designed a unique latitudinal soil translocation experiment to investigate the relationship between the bacterial and fungal community stability and the responses of soil C molecular structure to climate warming. The detailed protocol for the experiment was the following: (1) Forty kilograms of topsoil (0–25 cm) was collected for each SOM. The latitude and longitude of the sampling sites and soil geochemical characteristics are shown in Tables S3 and S4. Detailed data can be found in Supplementary Data 1. (2) The soil was homogenized using a 2 mm sieve and filled with sterilized PVC tubes. The PVC tube was 5 cm in diameter at the bottom and 31 cm in height. Each tube was filled with a 25 cm-high soil column, which corresponded to approximately 1 kg of soil. The bottom of the pipe was filled with 1 cm quartz sand, and a 5 cm space was left at the top. (3) From October to November 2015, 90 PVC pipes containing soil (5 SOM gradients × 3 replicates × 6 climatic conditions) were transported to six ecological research stations with different geoclimatic conditions and SOM contents, and 15 PVC pipes were placed in each station. Once the experiment was set up, the weeds growing in each PVC pipe were manually removed every 2–3 weeks to avoid the impact of plants.The six ecological research stations were the Hailun Agricultural Ecological Experimental Station (HL, N 47°27′, E 126°55′) in Heilongjiang Province, Shenyang Agriculture Ecological Experimental Station (SY, N 41°49′, E 123°33′) in Liaoning Province, Fengqiu Agricultural Ecological Experimental Station (FQ, N 35°03′, E 114°23′) in Henan Province, Changshu Agricultural Ecological Experimental Station (CS, N 31°41′, E 120°41′) in Jiangsu Province, Yingtan Red Soil Ecological Experiment Station (YT, N 28°12′, E 116°55′) in Jiangxi Province and Guangzhou National Agricultural Science and Technology Park (GZ, N 23°23′, E 113°27′) in Guangdong Province. The MAT and MAP at the six ecological research stations ranged from 1.5 to 21.9 °C and from 550 to 1750 mm from north to south, respectively. Details of their climatic conditions (e.g., climatic types) are shown in Table S5. All tubes were removed from each station after 1 year.The soil samples were stored on dry ice and rapidly transported back to the laboratory. The soil pH was measured by the potentiometric method. Nitrate (NO3−-N) and ammonium nitrogen (NH4+-N) were measured by the Kjeldahl method. DOC was measured using a total organic carbon analyzer (Shimadzu Corporation, Kyoto, Japan). SOC was determined by wet digestion using the potassium dichromate method53. Microbial biomass C (MBC) was measured by the chloroform fumigation-incubation method54. All geochemical attributes are shown in Table S4.Solid-state 13C NMR analysis of soil C molecular groupsSolid-state 13C NMR spectroscopy analysis was performed to determine the molecular structure of SOC. A Bruker-Avance-iii-300 spectrometer was used at a frequency of 75 MHz (300 MHz 1H). Before the examination, the soil samples were pretreated with hydrofluoric acid to eliminate the interference of Fe3+ and Mn2+ ions in the soil. Specifically, 5 g of air-dried soil was weighed in a 100 ml centrifuge tube with 50 ml of hydrofluoric acid solution (10% v/v) and shaken for 1 h. The supernatant was then removed by centrifugation at 3000 rpm for 10 min. The residues were washed eight times with a hydrofluoric acid solution (10%) with ultrasonication. The oscillation program consisted of the following: four × 1 h, three × 12 h, and one × 24 h. The soil samples were washed with distilled water four times to remove the residual hydrofluoric acid. The above-mentioned treated soil samples were dried in an oven at 40 °C, ground and passed through a 60-mesh sieve for NMR measurements.The soil samples were then subjected to solid-state magic-angle rotation-NMR measurements (AVANCE II 300 MH) using a 7 mm CPMAS probe with an observed frequency of 100.5 MHz, an MAS rotation frequency of 5000 Hz, a contact time of 2 s, and a cycle delay time of 2.5 s. The external standard material for the chemical shift was hexamethyl benzene (HMB, methyl 17.33 mg kg−1). The spectra were quantified by subdividing them into the following chemical shift regions55: 0–45 ppm (alkyl), 45–60 ppm (N-alkyl and methoxyl), 60–110 ppm (O-alkyl), 110–140 ppm (aryl), 140–160 ppm (O-aryl), 160–185 ppm (carboxy), and 185–230 ppm (carbonyl) (Fig. 3a). We classified O-alkyl, O-aryl, and carboxy C as labile C and alkyl, N-alkyl/methoxyl, and aryl C were classified as recalcitrant C.Soil microbial C metabolic profilesThe soil microbial C metabolic capacities were measured with BIOLOG 96-well Eco-Microplates (Biolog Inc., USA) using 31 different C sources and three replicates in each microplate. These C sources included carbohydrates, carboxylic acids, polymers, amino acids, amines, and phenolic acids (Table S2). Carbohydrates, amino acids, and carboxylic acids are generally considered labile C sources, amines and phenolic acid compounds are relatively resistant C sources, and polymers are recalcitrant C. The diverse nature of these C sources allowed us to identify differences in the capacity of microbes to degrade different C sources56. Soil microbes were extracted as follows: (1) Five grams of soil (dry weight equivalent) was incubated at 25 °C for 24 h, and 45 ml of sterile 0.85% (w/v) sodium chloride solution was added57. (2) At room temperature (25 °C), the mixture was shaken at 200 rpm for 30 min and allowed to stand for 15 min. (3) Subsequently, 0.1 ml of the supernatant was collected and diluted to 100 ml with sterile sodium chloride solution. (4) Soil suspensions were dispensed into each of the 93 wells (150 μl per well), and the plates were then incubated at 25 °C in the dark for 14 days. The optical density (OD, reflecting C utilization) of each well was read at 590 nm (color development) every 12 h. The normalized OD of different C sources was calculated as the OD of the well that contained the C source minus the OD of the well that contained sterile sodium chloride solution (control well). The normalized OD at a single time point (228 h) was used for the posterior analysis when it reached the asymptote.DNA extraction, PCR amplification, and sequencingDNA was extracted from all 90 soil samples. Briefly, well-mixed soil samples (0.6 g) were analyzed using the Power Soil DNA Isolation Kit (MoBio Laboratories, Inc., Carlsbad, CA, USA) following the manufacturer’s instructions. The quality of the DNA extracts was determined by spectrophotometry (OD-1000+, OneDrop Technologies, China). The DNA extracts were considered of sufficient quality if the ratio of OD260 to OD280 (optical density, OD) and the ratio of OD260 to OD230 were approximately 1.8. All eligible DNA samples were stored at −80 °C.Taxonomic profiling of the soil bacterial and fungal communities was performed using an Illumina® HiSeq Benchtop Sequencer. PCR amplification was performed using an ABI GeneAmp® 9700 (ABI, Foster City, CA, USA) with a 20 μl reaction system containing 4 μl of 5× FastPfu Buffer, 0.8 μl of each primer (5 μM), 2 μl of 2.5 mM dNTPs, 2 μl of template DNA, and 0.4 μl of FastPfu Polymerase. For bacterial analysis, the forward the primer 515F (GTGCCAGCMGCCGCGG) and the reverse primer 907R (CCGTCAATTCMTTTRAGTTT) were used to amplify the bacteria-specific V4-V5 hypervariable region of the 16S rRNA gene58. For fungal analysis, the internal transcribed spacer 1 (ITS1) region of the ribosomal RNA gene was amplified with primers ITS1-1737F (GGAAGTAAAAGTCGTAACAAGG) and ITS2-2043R (GCTGCGTTCTTCATCGATGC)59. The PCR protocol for bacteria consisted of an initial predenaturation step of 95 °C for 2 min, 35 cycles of 20 s at 94 °C, 40 s at 55 °C and 1 min at 72 °C, and a final 10 min extension at 72 °C. The PCR protocol for fungi consisted of an initial predenaturation step of 95 °C for 3 min, 35 cycles of 30 s at 95 °C, 30 s at 59.3 °C, and 45 s at 72 °C and a final 10 min extension at 72 °C.Each sample was independently amplified three times. Following amplification, 2 μl of each of the PCR products was checked by agarose gel (2.0%) electrophoresis, and all the PCR products from the same sample were then pooled together. The pooled mixture was purified using the Agencourt AMPure XP Kit (Beckman Coulter, CA, USA). The purified products were indexed in the 16S and ITS libraries. The quality of these libraries was assessed using Qubit@2.0 Fluorometer (Thermo Scientific) and Agilent Bioanalyzer 2100 systems. These pooled libraries (16S and ITS) were subsequently sequenced with an Illumina HiSeq 2500 Sequencer to generate 2 × 250 bp paired-end reads at the Center for Genetic & Genomic Analysis, Genesky Biotechnologies Inc., Shanghai, China.The raw reads were quality filtered and merged as follows: (1) TrimGalore was used for truncation of the raw reads at any site with an average quality score  5%) soils, changes in the C metabolic capacity of microbes under elevated temperatures were characterized using the ratio of the OD of microbes measured in the translocated soils to the OD of microbes in the in situ HL soil. A ratio greater than 1 indicates that translocation warming increases the C metabolism of microbes.Mantel and partial Mantel analysisA previous study showed that partial Mantel analysis is a robust method for evaluating the relationship among three variables65. This approach can control the z-axis and assess only the relationship between the x- and y-axes, avoiding the interaction between the z- and x-axes on the y-axis. In this study, Mantel analysis was employed to assess the relationships between the stability of the bacterial and fungal communities and C metabolic capacity. Stability refers primarily to the ability of the microbial community to resist translocation warming66. A higher similarity between the microbial communities in translocated soil compared with that in the in situ HL area indicates that the community is more resistant to translocation-related warming and that the microbial community is more stable.Calculation of the microbial β-diversityBray-Curtis and Euclidean dissimilarity metrics were calculated to estimate the bacterial and fungal taxonomic dissimilarity (β-diversity) and environmental dissimilarity (e.g., latitude, MAT, and MAP), respectively, using the vegan package (version 2.5–6) in the R statistical program (version 4.0.2, https://www.r-project.org/)67. Corresponding to the 45 C metabolism ratios in soils with the same OM content, the β-diversity values of bacteria and fungi were selected to analyze the relationship between the community similarity (1-β-diversity) of bacteria and fungi and changes in microbial C metabolism.Impact of the SOM content and climate change on changes in microbial communitiesThe distribution patterns of the bacterial and fungal communities under different SOM gradients and climatic regimes were determined through nonmetric multidimensional scaling (NMDS)68. To quantitatively compare the effects of the SOM gradient and climatic regimes on the bacterial and fungal community composition, three nonparametric multivariate statistical analyses were used in this study: nonparametric multivariate analysis of variance (Adonis), analysis of similarity (ANOSIM), and multiple response permutation procedure (MRPP)69. The linear fit between environmental dissimilarity and microbial β-diversity was analyzed using the lm function in R. A significant difference in the bacterial and fungal β-diversity among different SOM contents was evaluated by Student’s paired t-test using the ggpubr (version 0.4.0) package70. RDA was performed to analyze the relationships of bacterial and fungal communities with various environmental factors (soil geochemical attributes and climatic conditions, such as MAP and MAT). In parallel, the Monte Carlo permutation test (999 permutations) was employed to determine whether the explanation of the microbial distribution by individual factors (e.g., pH, SOC, and TN) was significant71.Construction of the structural equation model and random forest modelA SEM was fitted to illustrate the direct or indirect effects of soil properties (e.g., pH, moisture, ammonia, and nitrate nitrogen), climate change (e.g., MAT and MAP), and bacterial and fungal β-diversity on soil C metabolic capacity72. Based on the Euclidean method, the changes in soil properties and climatic conditions of five translocated sites compared with those in the in situ HL site were calculated. A total of 45 ratios were obtained for each OM content. Corresponding to the 45 ratios in soils with the same OM content, the β-diversity values of bacteria and fungi were selected. The model construction process was mainly divided into three steps. In brief, these steps include the establishment of an a priori model, data normality detection, and an overall goodness-of-fit test. The prior model was constructed based on a literature review and our knowledge. For the variables that did not conform to the normal distribution, we performed logarithmic transformation. Here, we used the χ2 test (the model was assumed to exhibit a good fit if p  > 0.05), the goodness-of-fit index (GFI; the model was assumed to show a good fit if GFI  > 0.9), the root mean square error of approximation (RMSEA; the model was assumed to exhibit a good fit if RMSEA  0.05)73 and the Bollen-Stine bootstrap test (the model was assumed to show a good fit if the bootstrap p  > 0.10) to test the overall goodness of fit of the SEM. All SEM analyses were conducted using IBM® SPSS® Amos 21.0 (AMOS, IBM, USA). Additionally, the importance of the metabolic capacity of different types of C on labile and recalcitrant C was assessed by random forest models using the randomForest package (version 4.6-14) in R74, and the model significance and amount of interpretation were evaluated using the rfUtilities package (version 2.1–5)75.Reporting summaryFurther information on research design is available in the Nature Research Reporting Summary linked to this article. More