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Experimental designConsortia of auxotrophic E. coli genotypes, which previously evolved an obligate mutualistic cooperation26, were used to determine how this type of interaction affects the ability of the participating individuals to respond to environmental selection pressures. To this end, two main experimental treatment groups were established. First, each of the two cooperative auxotrophs was grown as amino acid-supplemented monoculture (i.e. tyrosine and tryptophan, 100 µM each). Second, both genotypes were cocultivated in the absence of amino acid supplementation. A treatment, in which monocultures were cultivated in the absence of amino acid supplementation was not included, because auxotrophic genotypes would not grow under these conditions. Also, an amino acid-supplemented coculture was not implemented in the experimental design, because competition between both auxotrophs was likely to result in a loss of one of the two genotypes (Supplementary Fig. 1). Moreover, previous experiments showed that amino acid supplementation does not completely abolish the mutualistic interaction. Hence, the experiment compared monocultures with externally provided amino acids (i.e. no mutualism) to cocultures, which could only grow when strains reciprocally exchanged amino acids (i.e. mutualism). Replicate populations of both treatment groups were serially propagated while being subject to a stepwise increasing concentration of one of four different antibiotics (i.e. ampicillin, kanamycin, chloramphenicol, and tetracycline) (Fig. 1). These four antibiotics differed in their mode of action. In this way, not just the effect of a single stressor was probed, but rather the ability of mutualistic consortia to adapt to environmental stress in general.Ancestral consortia differ in their growth levels and susceptibility to environmental stressBefore the actual evolution experiment was performed, both growth levels and susceptibility to environmental stress was determined in the ancestral consortia. Comparing the maximum growth rate and densities populations achieved after 72 h revealed that unsupplemented cocultures grew significantly slower (Benjamini–Hochberg correction: P  More

Significance tests of straw return methods, potassium fertilization levels and their interactionsAnalysis of variance (ANOVA) results showed that straw return methods and potassium fertilization levels had significant effects on maize photosynthesis, dry matter and yield from 2018 to 2020 (Table 3). Significant interactions between straw return methods and potassium fertilization levels were only found on Pn of 2018 and 2020, and Tr of 2018–2020. Through the comparison of three-year F-values, it could be found that the effect of potassium fertilization levels on maize photosynthesis, dry matter and yield was greater than that of straw return methods.Table 3 Significance of the effects of straw return methods, potassium fertilization levels and their interactions on maize growth and yield using ANOVA.Full size tableEffects of straw return and potassium fertilizer on photosynthesis of maizeThe straw return methods and potassium fertilization levels significantly influenced (p ≤ 0.05) the maize photosynthesis compared to control (CK), resulting in Pn, Gs and Tr values that were higher than those of CK, and Ci value that was lower than that of CK.Straw return and potassium supply increased Pn, Gs and Tr. From 2018 to 2020, compared with CK, Pn increased by 1.70–4.09 under SFK0, 2.65–5.77 under SFK30, 5.21–8.48 under SFK45, 7.31–11.44 under SFK60, 0.63–3.20 under FGK0, 2.50–5.11 under FGK30, 3.60–5.79 under FGK45, and 3.97–7.47 μmol·m-2·s-1 under FGK60 (Fig. 1a). Gs increased by 0.60–0.90 under SFK0, 0.10–0.13 under SFK30, 0.18,-0.19 under SFK45, 0.20–0.22 under SFK60, 0.02–0.06 under FGK0, 0.08–0.09 under FGK30, 0.13–0.17 under FGK45, and 0.15–0.19 mmol·m-2·s-1 under FGK60 (Fig. 1b). Tr increased by 0.55–0.87 under SFK0, 1.02–1.30 under SFK30, 1.51–1.67 under SFK45, 1.74–1.99 under SFK60, 0.49–0.71 under FGK0, 0.86–1.13 under FGK30, 1.12–1.38 under FGK45, and 1.27–1.47 mmol·m−2·s−1 under FGK60 (Fig. 1c).Figure 1Effects of straw return methods and potassium fertilization levels on maize photosynthesis.Full size imageStraw return and potassium supply decreased Ci. From 2018 to 2020, compared with CK, Ci decreased by 5.43–8.92 under SFK0, 10.59–14.05 under SFK30, 19.04–21.21 under SFK45, 21.77–23.81 under SFK60, 2.26–6.52 under FGK0, 8.59–12.07 under FGK30, 12.93–16.15 under FGK45, and 17.81–19.46 μmol·mol-−1 under FGK60 (Fig. 1d).Comprehensive analysis showed that Pn, Gs, Tr increased and Ci decreased significantly after the treatment of SF under the same potassium supply. Under the same straw return method, Pn, Gs and Tr values increased significantly with the potassium fertilization levels, while Ci decreased. The effects of straw return and potassium fertilizer on maize photosynthesis increased gradually from year to year.Effects of straw return and potassium fertilizer on dry matter of maizeWe can see from Fig. 2, the straw return methods and potassium fertilization levels significantly increased (p ≤ 0.05) the maize dry matter accumulation. Compared with CK, under the treatments of SFK0, SFK30, SFK45, SFK60, FGK0, FGK30, FGK45 and FGK60, the dry matter of R1 and R6 stage increased by 1454.45, 2288.75, 3982.85, 4961.45, 1042.96, 1744.54, 2890.65, 3408.39 and 2152.43, 4433.55, 6726.72, 8051.51, 1195.76, 3337.79, 5121.77, 6247.56 kg/ha in 2018; the dry matter increased by 1812.69, 2959.44, 4370.19, 5615.94, 1545.06, 2238.06, 3421.11, 4028.64 and 2588.52, 5319.60, 7500.74, 8912.64, 1649.67, 3832.46, 6065.90, 6864.33 kg/ha in 2019; the dry matter increased by 2535.39, 3612.35, 5544.00, 6720.12, 2474.18,2827.94, 4749.86, 4769.66 and 3235.18, 5798.75, 8577.48, 10,071.83, 2515.75, 4386.39, 7256.61, 7536.91 kg/ha in 2020.Figure 2Effects of straw return methods and potassium fertilization levels on maize dry matter. Values followed by different letters in the same year indicated indicate statistical significance at α = 0.05 under different treatments. The same below.Full size imageIn short, under the same straw return method, the increase of maize dry matter from R1 to R6 improved significantly with the potassium level, potassium fertilizer could improve the maize dry matter accumulation ability. The maize dry matter of R1 to R6 increased significantly after the treatment of SF compared to FG under the same potassium supply. The promotion effect of straw return and potassium fertilizer on maize dry matter increased from year to year.Effects of straw return and potassium fertilizer on maize yieldThe straw return methods and potassium fertilization levels significantly influenced (p ≤ 0.05) the maize yield compared to CK, resulting in maize yield values that were higher than those of CK. Straw return and potassium supply increased maize yield. From 2018 to 2020, compared with CK, maize yield increased by 9.73–10.32% under SFK0, 15.68–17.47% under SFK30, 24.02–25.58% under SFK45, 24.46–25.76% under SFK60, 5.79–7.83% under FGK0, 13.51–13.72% under FGK30, 18.64–19.01% under FGK45, and 21.19–21.69% under FGK60 (Fig. 3).Figure 3Effects of straw return methods and potassium fertilization levels on maize yield.Full size imageThe maize yield among treatments was as follows: SFK60  > SFK45  > FGK60  > FGK45  > SFK30  > FGK30  > SFK0  > FGK0  > CK. Compared to FG, the effect of SF on maize yield was more obvious. The maize yield increased significantly with the potassium fertilization levels under the potassium fertilization levels of 0–60 kg/ha in this test. The treatment of SFK60 recorded the highest average yield in the three-year test, which was 14,744.39 kg/ha. The maize yield in different planting years showed as follows: 2020  > 2019  > 2018, which indicated that the promotion effect of straw return and potassium fertilizer on maize yield increased from year to year.Correlation analysis of photosynthesis, dry matter accumulation and yield of maizePn, Gs, Tr and Ci were significantly correlated with dry matter accumulation. Pn, Gs and Tr were positively correlated with dry matter, while Ci was negatively correlated with the dry matter (Table 4). The results showed that the increase of Pn, Gs, Tr and the decrease of Ci could significantly improve maize dry matter. Dry matter was positively correlated with maize yield, indicating that the increase of dry matter accumulation could significantly improve maize yield. The increase of Pn, Gs, Tr and dry matter accumulation, as well as the decrease of Ci, could significantly increase maize yield.Table 4 Correlation analysis of photosynthesis, dry matter accumulation and yield of maize under two straw return methods.Full size tableUnder the method of SF, the correlation coefficients of Pn, Gs, Tr, dry matter at R1 stage, dry matter at R6 stage and Ci with yield were 0.862, 0.988, 0.962, 0.948, 0.971 and −0.978; the correlation coefficients were 0.838,0.975,0.970,0.930,0.979 and −0.973 under the method of FG. The results showed that, under the method of SF, the correlation coefficients between dry matter of R1 stage, Pn, Gs, Ci with yield were higher than that under the method of FG, which indicated that SF could promote the correlation between the dry matter of R1 stage, Pn, Gs, Ci with yield. Under the method of FG, the correlation coefficients between the dry matters of R6 stage, Tr with yield were higher than that under the method of SF, which indicated that FG could promote the correlation between the dry matter of R6 stage, Tr with yield.Effects of straw return and potassium fertilizer on the profit of maize plantingGross income is an important economic index that determines the profit or benefit that a farmer can obtain. On the other hand, net return reflects the actual income of the farmer. According to the average selling price of maize (1 yuan/kg) from 2018 to 2020, the net income of maize planting of different treatments was as follows: SFK45  > SFK60  > FGK60  > FGK45  > SFK30  > FGK30  > SFK0  > FGK0  > CK (Table 5). Compared to CK. the average net profit of maize planting in the three-year test increased by 421.26, 1049.07, 2014.82, 1980.44, 313.58, 1035.34, 1587.44, 1828.69 yuan/ha between the treatments of SFK0, SFK30, SFK45, SFK60, FGK0, FGK30, FGK45 and FGK60. Straw return and potassium supply increased the net profit of maize planting. The net profit of maize planting increased significantly after SF compared to FG under the same potassium supply. The treatment of SFK45 reached the maximum profit of maize planting, which was 2014.82 yuan/ha.Table 5 Effects of straw return methods and potassium fertilization levels on the profit of maize planting.Full size table More

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