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Komercijalna banka bircaninova kontakt torrent 26.06.2020

arctic paper grycksbo kontakt torrent

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Temperature is likely to be an important control on P bioavailability, although effects may differ across contrasting plant communities with different soil properties. We used an elevational gradient in northern Sweden that included both heath and meadow vegetation types at all elevations to study the effects of temperature, soil P sorption capacity and oxalate-extractable aluminium Al ox and iron Fe ox on the concentration of different soil P fractions.

We hypothesized that the concentration of labile P fractions would decrease with increasing elevation and thus declining temperature , but would be lower in meadow than in heath, given that N to P ratios in meadow foliage are higher. As expected, labile P in the form of Resin-P declined sharply with elevation for both vegetation types.

Meadow soils did not have lower concentrations of Resin-P than heath soils, but they did have 2—fold and 1. Additionally, Resin-P expressed as a proportion of total soil P for the meadow was on average half that in the heath. Declining Resin-P concentrations with elevation were best explained by an associated 2.

In contrast, the lower P availability in meadow relative to heath soils may be associated with impaired organic P mineralization, as indicated by a higher accumulation of organic P and P sorption capacity. Our results indicate that predicted temperature increases in the arctic over the next century may influence P availability and biogeochemistry, with consequences for key ecosystem processes limited by P, such as primary productivity.

This is an open-access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: The authors have declared that no competing interests exist. Phosphorus P is an important macronutrient in subarctic tundra, where it is often co-limiting with nitrogen N , and is sometimes the main limiting nutrient for plant growth [1] — [3].

Most soil P in the surface of tundra soils is organic [1] , [4] , [6] and is dominated by highly labile compounds [7]. Temperature is one of the main controls of organic matter decomposition in the arctic [8] — [10] , meaning it is likely to constrain organic P mineralization and the supply of available P for plants. Likewise, an increase in foliar and litter P concentrations, together with a decrease in foliar and litter N to P ratios indicative of greater relative P availability , have been observed with decreasing elevation thus increasing temperature in subarctic tundra [13].

Current knowledge on the distribution of different P forms in tundra landscapes and how they may be affected by temperature is limited, despite this information being crucial for understanding future temperature effects on bioavailable P. Elevational gradients are powerful tools for studying how temperature and associated climatic factors, which shift with elevation, influence ecosystems properties and processes [17] — [21].

As such, an increasing number of studies in a wide range of ecosystems have used elevational gradients to study how temperature affects ecological processes [22] — [24] including in the subarctic [13] , [25] — [28]. Elevational gradients also provide excellent opportunities for exploring the impacts of temperature on the availability of soil P over larger spatial scales and timeframes than what is possible through conventional experiments [19].

There are, to our knowledge, no studies available on the responses of different P pools to elevation in subarctic or arctic tundra. These landscapes are biogeochemically heterogeneous as a consequence of spatial variation in topography and plant community structure [29] , [30]. This results in high spatial variation in P availability [6] , [25] and in the concentrations of Al and Fe [6] , [31] , which influence P availability [32] , [33].

The Fennoscandian tundra therefore consists of a mosaic of highly contrasting vegetation types. Specifically, heath vegetation occurs on soils with low pH and N availability and is dominated by slow-growing dwarf-shrubs, while meadow vegetation grows on soils that are more N-rich and is dominated by faster-growing herbaceous species [25] , [29] , [30]. The biogeochemistry of P differs between the two vegetation types [6] and soil phosphate concentrations and plant foliar N to P ratios suggest that the relative importance of P versus N limitation is greater in meadow than in heath [6] , [13] , [25].

As such, obtaining a representative picture of how P biogeochemistry varies in the subarctic requires explicit recognition of both vegetation types. In this study, we used a well-established elevational gradient [13] , [25] , [26] , [34] in which both heath and meadow vegetation types occur at all elevations, to study the effects of elevation-associated variation in temperature on P availability and biogeochemistry in a subarctic ecosystem. We also used this gradient to examine whether previously reported changes in foliar and litter P contents and N to P ratios with elevation are matched by shifts in the concentration of P fractions of different lability.

To determine P fractions we used the Hedley fractionation method [35] — [38] , an approach widely used to determine landscape-level variation in P availability and dynamics [6] , [39] , [40] — [42]. Additionally, we examined if other known drivers of P availability, such as soil P sorption capacity and Al and Fe concentration, influence the distribution of P fractions across the gradient for both the heath and meadow vegetation.

Specifically, we tested the hypotheses that 1 The concentrations of labile P fractions decline with elevation and therefore temperature regardless of vegetation type, 2 Across all elevations, meadow soils have consistently lower concentrations of labile P than heath soils, together with higher concentrations of Al and Fe and higher soil P sorption capacity.

By addressing these hypotheses we aim to better understand how temperature changes, such as those that are expected through climate warming, may affect P availability across two dominant vegetation types in subarctic tundra ecosystems. This study was carried out across an elevational gradient ranging from to m above sea level a. Figure 1 shows a map with the location of the study site and the elevational gradient.

No part of this gradient is located within national reserves and the land is public and not government protected. We confirm that all national and international rules were observed during the field work. This investigation did not involve measurements on animals or humans. The soils collected for this research were sampled at very small spatial scales and thus had negligible effects on ecosystem functioning.

We have no commercial interests or conflicts of interest in performing this work. Filled black circles indicate each of the six study sites, ranging in elevation from to The mean annual precipitation in the area, measured at the Abisko Scientific Research Station, was mm for the period —, with the highest mean monthly precipitation in July 51 mm and the lowest in April 12 mm [43]. The treeline in this area is formed by Betula pubescens spp. Two types of vegetation, heath and meadow, grow in a mosaic across the study area and co-occur on all elevations, with the meadow generally found in shallow depressions.

The heath is dominated by ericaceous dwarf-shrubs such as Vaccinium vitis-idaea , V. The meadow vegetation is dominated by graminoids such as Deschampsia flexuosa and Anthoxanthum alpinum , herbs such as Saussurea alpina , Viola biflora and Solidago virgaurea , and sedges, notably Carex bigelowii [25].

The bedrock is comprised of salic igneous rocks and quartic and phyllitic hard schists. The soils are podzols at lower elevations and cryosols at higher elevations. For more details on the study system see Table S1 and [13] , [25].

To minimize pseudoreplication within each elevation, the average distance between each plot and the next nearest plot was c. Because the microtopography, hydrology, and soil fertility of these communities is highly spatially heterogeneous over short distances i. Plots at the m elevation site were located in open birch forest, plots at m were situated immediately above the forest line, and plots from m to m were devoid of trees [25].

Monthly mean air temperatures during August at m, m and m at the study site were The daily mean temperature across the elevational gradient during the growing season of is given in Figure S1 ; similar data for the previous year is also given in Figure S2 [25].

For each plot, a minimum of four 4. Within each plot, the cores were sieved 2 mm mesh in the field to homogenize the samples, and combined to yield a single bulked sample per plot. Samples were sealed in polyethylene bags and transported to the laboratory on the same day as sampling.

In order to characterize soil P composition, we performed a five-step sequential extraction [38] with some modifications [44] , [45] , outlined in Figure 2. This method was chosen because it provides a direct estimate of the lability of different operationally-defined P pools, and is the most commonly used method to investigate differences in P availability and dynamics in natural soils [35] — [37] , [46].

The resin was transferred to a bottle and eluated on a shaker 1 h, rpm with 40 mL NaCl. In Step 2, the soil remaining from Step 1 was combined with mL 0. In Step 3, the soil remaining from Step 2 was extracted with 0.

In Step 4, the soil remaining from Step 3 was combined with 1. In the final and fifth step, the soil remaining from Step 4 was washed with mL of deionised water by shaking for 1 h, centrifuging, and discarding the supernatant, after which the soil was set to air dry at room temperature.

Total labile P consists of resin-extractable and bicarbonate-extractable P [48]. Resin-P is well correlated with P uptake by plants [49] , [50] , has a rapid turnover and high bioavailability, and consists of P in a form that can exchange freely between the solid phase of the soil and the soil solution [35] , [51].

Bicarbonate-P i Bic-P i has similar sources to resin-P, turns over fast and is also bioavailable in the short-term [35] , while bicarbonate-P o Bic-P o is easily mineralizable and supplies plant-available P. HCl-extractable P represents calcium-bound P i and is often taken to represent P associated with primary minerals; it is also considered to be more stable [48].

In order to estimate the relative P sorption capacity of each soil sample from each plot, we used a single point P sorption method [52]. For each sample, 2 g dry weight of soil were weighed into each of two 60 mL bottles respectively, and 40 mL of mM KCl was added to each bottle. For one of the two bottles, 1. The suspensions were shaken for 24 h, filtered Munktell 00H filter paper, pore size approx. The amount of sorbed P was estimated as the difference between the phosphate concentration in spiked and unspiked samples.

Thus, a high index indicates a high P sorption capacity. The concentration of organic P was calculated as the difference between total and inorganic P. Phosphorus in the digests was analysed as above. Aluminium Al and iron Fe concentrations in the soil samples were determined following extraction by 0. A subsample of 0. The oxalate extractant is assumed to release exchangeable Al and Fe and dissolve non-crystalline and poorly crystalline oxides of Al and Fe i.

We chose ANOVA because it is the most powerful way of detecting significant responses to the underlying gradient even when these responses are not unidirectional or simple [56]. To further explore the effects of elevation within vegetation types, one-way ANOVA testing for the effect of elevation was performed separately for both heath and meadow.

Where significant effects of elevation were found, data were further analyzed for differences among means using Tukey's honestly significant difference h. Tukey's h. In order to account for potential effects of co-variation of soil P sorption capacity with elevation on labile P concentrations, we divided the concentration of each of the three labile components i. We then performed separate one-way ANOVA testing for the effect of elevation on the transformed data, followed by Tukey's h.

When required, data were transformed to conform to the assumptions of parametric tests. Total labile P i. The concentrations of the remaining fractions ranged from 0. The most unidirectional effect of elevation was found for Resin-P, for which the highest concentrations were recorded at the lowest elevation and the lowest concentrations at the highest elevation, for both vegetation types Figure 3 , Table 1.

Resin-P concentrations at the highest elevation were 7—fold and 11—fold lower than at the lowest elevation for heath and meadow, respectively. Total labile P trends mirrored those of Resin P, concentrations at the highest elevation m in heath were less than one fifth of those recorded at the m lowest elevation and m sites. In meadow, Total labile P concentrations at the highest elevation were less than one third of those recorded at the lowest elevation.

After concentrations were divided by the sorption index, the unidirectional elevational trend remained for Resin-P and the non-unidirectional effect remained for Bic-P o ; further, a significant effect of elevation emerged for Bic-P i , through it being highest at the lowest elevation for the meadow Figure S3.

Finally, Resin-P and Total labile P were significantly positively related with temperature in both the heath and meadow vegetation Table S3 , as was the case for pH in the heath. Soils were collected in subarctic heath and meadow vegetation along an elevational gradient — m in Abisko, Sweden.

Note the difference in y-axis scales. Soils were collected in subarctic heath and meadow vegetation types along an elevational gradient — m in Abisko, Sweden. The concentrations of all P fractions except Resin and Total labile P were significantly different between meadow and heath, and of these all were highest in the meadow except for Bic-P o Table 1 , Figure 3. After concentrations were divided by the sorption index, Bic-P i and Bic-P o remained highest in the meadow and heath, respectively, and Resin-P was also highest in the heath Figure S3.

There was an interactive effect between vegetation type and elevation for all P fractions except Bic-P i and HCl-P, meaning that soil P composition responds differently to changes in elevation depending on vegetation type Table 1 , Figure 3. The concentrations of labile P fractions were hypothesized to decline with increasing elevation and associated declines in temperature, regardless of vegetation type.

We consider Resin-P, Bic-P i and Bic-P o to represent the most labile P fractions [35] ; Resin-P is well correlated with plant P uptake [49] , [59] , and is considered to be the most highly bioavailable fraction [35] , [51]. Further, Bic-P i is considered to derive from similar sources to Resin-P, while Bic-P o to be easily mineralized [35] , [48].

These three fractions are discussed separately given the differences in their concentration and dynamics observed in this study. Partially in line with our predictions, Resin-P concentration declined with elevation in both heath and meadow. Concentrations of Bic-P i and P o did not show any unidirectional trends with elevation, but given that they occurred in much lower concentrations, the overall trend is still one of declining bioavailable P with increasing elevation.

A number of factors could explain the observed decrease in Resin-P with elevation. Organic P as NaOH-extractable and Residual P is the dominant form of P in these humus soils and enzymatic hydrolysis of organic P is a likely driver for the release of bioavailable inorganic P, as has been shown for Alaskan tundra [1].

Temperature is the strongest driver of soil enzyme activity in the subarctic [63] and warming experiments show that even a relatively small increase in temperature 1. The average air temperature difference across our elevational gradient during the vegetation growing season is about 2. This suggests that the decline in temperature associated with increasing elevation is an important driver of the elevational decline in Resin-P observed and is consistent with our hypothesis.

However, further information on organic P mineralization processes, for example using mineralization and soil enzyme studies together with 31 P nuclear magnetic resonance NMR spectroscopy to characterize soil organic P [65] , [66] , would be useful to increase our mechanistic understanding of the trends observed here.

Our results are in line with the few studies that have measured labile inorganic P concentrations at different elevations in subarctic tundra, as they report lower concentrations of 1. Our results are also consistent with [13] who showed a decrease in foliar P concentrations and an increase in foliar N to P ratios with elevation independent of vegetation type. Decreasing organic P mineralization with increasing elevation should lead to the accumulation of soil organic P, which was the case for NaOH-P o in heath Figure 3.

However, a greater accumulation of soil organic P with elevation may have been negated to some extent by declining primary productivity, as is observed with increasing elevation both in the vicinity of the study area [28] and globally [19]. Temperature can affect P availability directly via effects on microbial mineralization [68] and soil process rates [69] but also indirectly by influencing factors that affect soil processes such as plant [70] and microbial [71] community composition.

Temperature variation across the elevational gradient studied here has previously been shown to be related to a range of variables including soil pH, total N and ammonium concentration, C to N ratio, vegetation density, plant and microbial community composition, and fungal to bacterial ratios [25]. As such, many of the soil and vegetation properties that vary along the elevational gradient are likely to represent indirect temperature controls on P availability.

Our interpretation is supported by many other studies that have used elevational gradients to understand how temperature affects ecological properties and processes [17] , [22] — [25] , [72] — [75]. To our knowledge, this is the first time that a Hedley P fractionation analysis has been carried out along a subarctic elevational gradient [6] and the two most comprehensive reviews on P fractionation lack data for these ecosystems [35] , [36].

These data are necessary to constrain soil P pools in terrestrial biogeochemical models, which are invaluable to understand the processes controlling P cycling and the role of P in driving terrestrial plant productivity [36]. Additionally, the highest Resin-P concentrations measured i. While the concentration of Total labile P sum of Resin P and Bicarbonate-extractable P is considered to be low in tundra relative to other ecosystems [1] , [4] — [6] , our results show that this may not necessarily hold when a wide range of elevations is considered.

The 3—5 fold variation in Total labile P concentrations that we found along this elevational gradient encompasses the whole range of Total labile P concentrations recently reported in a world synthesis of Hedley P fractionation studies in natural ecosystems that spans 11 soil orders [36]. Our results further suggest that because small changes in elevation and thus temperature were associated with large changes in available P, increases in temperature according to current climate change predictions [14] — [16] may have a significant impact on future P availability in arctic tundra.

We hypothesized that the concentration of labile P fractions Resin-P, Bic-P i and Bic-P o would be lower in soils under meadow than heath vegetation, concomitant with higher Al and Fe concentrations and soil P sorption capacity. Nevertheless, the concentration of Bic-P i showed the opposite pattern and the concentration of Resin-P, the largest labile P pool, was not significantly different between vegetation types.

No correlations have been reported for other organic soils that have Al ox and Fe ox concentrations more within the range of what we measured here [6] , [40]. Taken together, our results suggest that the Al and Fe concentrations in meadow soils are insufficient to exert a strong control on the concentration of labile inorganic P. This is suggested by the fact that concentrations of NaOH-P o and Residual P which in our soils is mostly organic were 2.

Some organic P compounds have a high affinity for Al and Fe oxides [79] — [81] , and soil organic P is often strongly positively related with Al and Fe concentration in organic soils, as seen in both this Table 3 and other studies [40] , [45] , [61] , [77] , [82] , [83]. Soil organic P is generally considered to be less prone to sorption than are labile inorganic P forms such as Resin-P [84] , but in some humus soils organic P is correlated with Al ox and Fe ox concentration while labile inorganic P is not [40] , which is consistent with our findings.

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