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The generalist accumulates extracellular nitriteWe first tested whether the generalist accumulates substantial extracellular nitrite under our experimental conditions, and thus creates a niche for the specialist. To accomplish this, we grew the generalist alone in bioreactors with anoxic ACS medium amended with 12 mM nitrate as the growth-limiting substrate and measured the extracellular concentrations of nitrate and nitrite over time. We performed these experiments at pH 6.5 (strong nitrite toxicity) and 7.5 (weak nitrite toxicity).We observed a substantial accumulation of extracellular nitrite regardless of the pH (Fig. 3A, B). When grown at pH 6.5 (strong nitrite toxicity), extracellular nitrite accumulated to a concentration comparable to the initial nitrate concentration (measured maximum extracellular nitrite concentration, 11.8 mM; measured initial nitrate concentration, 12.0 mM) and was subsequently consumed to below the detection limit (Fig. 3A). When grown at pH 7.5 (weak nitrite toxicity), extracellular nitrite again accumulated to a concentration comparable to the initial nitrate concentration (measured maximum extracellular nitrite concentration, 11.7 mM; measured initial nitrate concentration, 12.9 mM) and was subsequently consumed to below the detection limit (Fig. 3B). During growth at pH 6.5, substantial nitrite consumption did not begin until a prolonged period of time after nitrate consumption was complete, resulting in a relatively long duration of nitrite availability (Fig. 3A). During growth at pH 7.5, in contrast, substantial nitrite consumption began immediately after nitrate consumption was complete, resulting in a relatively short duration of nitrite availability (Fig. 3B). The longer duration of nitrite availability at pH 6.5 indicates that the duration of the niche created by the generalist for the specialist depends on pH.Fig. 3: Growth and nitrogen oxide dynamics of the generalist in batch culture.We grew the generalist alone in a bioreactor at A pH 7.5 (weak nitrite toxicity) or B pH 6.5 (strong nitrite toxicity) under anoxic conditions with nitrate as the growth-limiting substrate. Blue squares are measured extracellular nitrate concentrations, yellow triangles are measured extracellular nitrite concentrations, and black circles are measured cell densities. We measured extracellular nitrate and nitrite concentrations with IC and cell densities with FC. C Measured durations of nitrite availability for the generalist growing in batch culture. We grew the generalist alone in 96-well microtiter plates under anoxic conditions with nitrate as the growth-limiting substrate. Open symbols are durations of nitrite availability at pH 6.5 and closed symbols are durations of nitrite availability at pH 7.5. Each symbol is an independent biological replicate.Full size imageTo routinely quantify the duration of nitrite availability, we grew the generalist alone with varying amounts of nitrate as the growth-limiting substrate. We then quantified the length of time from when the growth rate with nitrate was maximum to when the growth rate with nitrite was maximum. This cell density-based proxy measure is valid because the growth of the generalist is directly linked to the consumption of nitrate and nitrite (Fig. 3A, B). The cell density of the generalist was initially linearly correlated with nitrate consumption at both pH 6.5 (strong nitrite toxicity) (two-sided Pearson correlation test; r = −0.96, p = 1.5 × 10–8, n = 15) (Fig. 3A) and 7.5 (weak nitrite toxicity) (two-sided Pearson correlation test; r = −1.00, p = 2.2 × 10–16, n = 30) (Fig. 3B). After nitrate was depleted, the cell density of the generalist became linearly correlated with nitrite consumption at both pH 6.5 (strong nitrite toxicity) (two-sided Pearson correlation test; r = −0.97, p = 3 × 10–4, n = 7) (Fig. 3A) and 7.5 (weak nitrite toxicity) (two-sided Pearson correlation test; r = −0.97, p = 6.8 × 10–10, n = 16) (Fig. 3B). We further validated our cell density-based approach by testing for concordance with our IC-based direct measures of the duration of nitrite availability. We observed a significant positive and linear relationship between the cell density- and IC-based measures (two-sided Pearson correlation test; r = 0.999, p = 0.023, n = 3) (linear regression model; slope = 1.19, intercept = −2.31, r2 = 0.99) (Supplementary Fig. S2), which further validates our cell density-based approach to routinely estimate the duration of nitrite availability.Using our cell density-based approach, we found that the duration of nitrite availability was significantly longer at pH 6.5 (strong nitrite toxicity) than at 7.5 (weak nitrite toxicity) regardless of the initial nitrate concentration (two-sample two-sided t-tests; Holm-adjusted p  0.92, Holm-adjusted p  0.6), and thus followed model predictions (Fig. 4A). However, when the specialist was initially rare (measured initial log rS/Gs of –3.19, –2.65, and –0.88), the relative abundances of the specialist continuously decreased between the third and twelfth transfers (Mann–Kendall trend tests; tau = –0.61 to –0.89, p  0 were dominated by phenotype C (dominant ancestral phenotype with a long time delay between nitrate and nitrite consumption), while generalist isolates from co-cultures with initial rS/Gs  More

Hydrological changes in high-latitude peatlandsWe observed three hydrological pathways, i.e., drying, wetting, and fluctuating, for both peatland clusters, non-permafrost and permafrost peatlands (Fig. 2). Approximately 54% of the studied peatlands have shifted towards drier surface conditions since 1800 CE and more intensively since 1900 CE (Fig. 2a, d), which is in line with the post-LIA warming. The overall change point to drier conditions was dated to ca. 1950 CE for non-permafrost sites and ca. 1890 CE for the permafrost compiled group. Approximately 32% of the studied peatlands have shifted towards wetter conditions (Fig. 2b, e). The overall shifting point to wetter conditions was dated to ca. 1995 CE for non-permafrost peatlands and to ca. 1990 CE for permafrost peatlands. Wetting has been especially intensive since 1900 CE for non-permafrost peatlands and since 1950 CE for permafrost peatlands. Interestingly, the data showed that in permafrost peatlands a significant dry shift always preceded a wet shift (Fig. 2e). Approximately 14% of the studied peatlands indicated no clear trend, with fluctuating hydrological conditions (Fig. 2c, f).Non-permafrost peatlands generally showed spatially extensive drying across the northern high latitudes, apart from northeastern Canada, where a wetting trend was more frequently observed. Permafrost peatlands, however, were more variable, with some drying, some wetting, and no overall coherent regional pattern was visible (Fig.1a, b). It should be noted that peatlands synthesized here have undergone little or no direct human impact, i.e., their surface hydrology was not significantly affected by human disturbances such as drainage, when compared to, for example, central European peatlands discussed in Swindles et al. (2019)5. This implies that climate and/or local topographical forcing are the predominant hydrological drivers in this study. The dataset is to some extent biased as there are more non-permafrost records from northeastern Canada but more permafrost records from northern Sweden and this might result in regional overestimation to either wetting or drying trends. Nevertheless, the pattern of diverse timing of the hydrological shifts between the individual coring points (Fig. 2) indicates the variability in sensitivity of different regions/peatlands to climate changes.Potential links to climate change and permafrost dynamicsThe comparison of the reconstructed water table and climate data suggests that climate, especially summer temperature, has played an important role in shaping the peatland water table (Fig. 1c–f). The pattern detected here for non-permafrost peatlands, an extensive drying, is comparable to that observed for central European peatlands5. In addition to direct climate forcing, a recent acceleration of peat accumulation might partly explain the drying trend by disconnecting the peatland surface from the water table17. However, for northeastern Canada a wetting trend has been observed more often, possibly regulated by the regional climate that shows clearly less warming in the focused period compared to other regions (Fig. 1c, d).Permafrost initiation in the past caused a peat surface uplift and is probably detected as a dry phase (Fig. 2d, e). Post-LIA warming-induced increase in evapotranspiration may have strengthened the surface drying which originally resulted from surface-uplift and probably mitigated the gradual wetting related to permafrost thawing11. The level of warming has varied among the regions. In some areas such as northeastern Canada temperature has increased less and, when combined with higher precipitation or higher effective moisture level, may have caused surface wetting in permafrost peatlands. This is a direct climate forcing rather than permafrost thawing, which is a consequence of climate warming, i.e., more indirect climate forcing. To date, it is yet challenging to estimate any one tipping point of warming that might trigger permafrost thawing, as the local conditions vary from bottom ground soil conditions to hydrology and vegetation. The consequent wetting or drying depends on evapotranspiration and ice richness etc., which further challenges the prediction of hydrological conditions of permafrost peatlands.The divergent three moisture patterns may occur in the same region and even in the same peatland, especially if the permafrost is present. This complex response pattern is well supported by the records from the Abisko region, Sweden, where replicated sampling was carried out, and captured different successional stages of local permafrost peatlands7,18. In contrast to permafrost peatlands, non-permafrost peatlands are more likely to experience a more consistent ecosystem response pattern19 as supported by the replicated records suggesting the same pattern happening simultaneously in several regions (Fig. 1a). The fluctuating pattern of many records reported here suggests that the past and recent climate has not yet caused a state change in hydrological conditions.Insights into carbon dynamics and future perspectiveGenerally, our results suggest that the recent climate warming has caused hydrological shifts in most high-latitude peatlands, highlighting its pronounced effect on shaping peatland moisture balance, and further on driving peatland C balance. It has been reported that a 1-cm water-table drawdown would increase 3.3–5.0 mg CO2 m−2 h−1 and decrease 2.2–3.6 mg CO2-eq m−2 h−1 (CH4) to the atmosphere, and the average sensitivity of CO2 and CH4 combined was 0.8–2.3 mg CO2-eq m−2 h−1 cm−1 according to a global scale analysis, including sites from high-latitudes3. However, it should be noted that the sensitivity of greenhouse gas fluxes to the magnitude of hydrological changes might vary among different regions and peatland types. It appears that most pan-Arctic peatlands are undergoing a drying trend, that may lead to a decreased C sink capacity3,19, if not compensated by increased C uptake from the atmosphere20.It is very likely that over the 21st century warming in high latitudes will continue to be more pronounced than the global average21. Precipitation is projected to increase, albeit with large regional variability. Also, extreme events with heavy rainfall and drought are becoming more frequent and intense22. It is estimated that about 20% of permafrost zone is experiencing accelerated and abrupt permafrost thaw that is likely causing wetting conditions4, while gradual permafrost thaw has been observed across the circumpolar regions23. Both an increase in precipitation and permafrost thaw might mitigate the drying pressure caused by warming and increased evapotranspiration. However, abrupt permafrost thaw in peatlands can result in a rapid (over years to decades) loss of C from the formerly frozen permafrost peat, causing these peatlands to be a net source of C to the atmosphere before post-thaw accumulation returns them to a net sink (centuries to millennia)12,13. The future C sink and source function of peatlands is a key element in contributing to climate change, but the observed divergent pathways of peatland hydrological successions further challenge the projections of high-latitude peatland C sink and source dynamics. Conversely, it clearly highlights the importance of climate forcing in peatland succession scenarios. Our study reveals that the response of high-latitude peatlands to changing climate conditions is complex. We detect variable ecohydrological trajectories, and in the future, these will determine the C sink capacity of northern peatlands. The observed patterns inevitably create challenges for the climate change modelling community. How to capture the highly heterogenic successional pathways of northern peatlands needs to be a key research focus. More

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