Icelandic grasslands as long-term C sinks under elevated organic N inputs

About 10% of the anthropogenic CO₂ emissions have been absorbed by northern terrestrial ecosystems during the past decades. It has been hypothesized that part of this increasing carbon (C) sink is caused by the alleviation of nitrogen (N) limitation by increasing anthropogenic N inputs. However, little is known about this N-dependent C sink. Here, we studied the effect of chronic seabird-derived N inputs (47-67 kg N ha⁻¹ year⁻¹) on the net soil organic C (SOC) storage rate of unmanaged Icelandic grasslands on the volcanic Vestmannaeyjar archipelago by using a stock change approach in combination with soil dating. We studied both early developmental (young) soils that had been receiving increased N inputs over a decadal timescale since an eruption in 1963, and well-developed soils, that had been receiving N inputs over a millennial timescale. For the latter, however, the effects on both decadal (topsoil; 40 years) and millennial (total soil profile; 1600 years) SOC storage could be studied, as the age of topsoil and the total soil profile could be determined from volcanic ash layers deposited in 1973 and 395 AD. We found that enhanced N availability-either from accumulation over time, or seabird derived-increased the net SOC storage rate. Under low N inputs, early developmental soils were weak decadal C sinks (0.018 ton SOC ha−1 year−1), but this increased quickly under ca. 30 years of elevated N inputs to 0.29 ton SOC ha⁻¹ year⁻¹, thereby equalling the decadal SOC storage rate of the unfertilized well-developed soils. Furthermore, for the well-developed soils, chronically elevated N inputs not only stimulated the decadal SOC storage rate in the topsoil, but also the total millennial SOC storage was consistently higher. Hence, our study suggests that Icelandic grasslands, if not disturbed, can remain C sinks for many centuries under current climatic conditions and that chronically elevated N inputs can induce a permanent strengthening of this sink

Temperature responses in a subarctic springtail from two geothermally warmed habitats

Common-garden experiments with populations sampled along natural thermal gradients help to reveal local adaptation, disentangle environmental and genetic effects, and ultimately predict, by analogy, future biotic responses to climate change. In this regard, geothermal habitats are useful model systems as they exhibit dramatic changes in soil temperature. The springtail Protaphorura pseudovanderdrifti has apparently coped with such local geothermal warming in Iceland, as this species occurs along a more than half-century-old geothermal gradient in a grassland and persists along a newly emerged temperature gradient in a previously non-geothermal planted spruce forest. We measured thermal reaction norms for development and walking speed and acute cold shock tolerance of P. pseudovanderdrifti originating from the grassland and forest geothermal gradients. Temperature-dependent juvenile development showed little variation among subpopulations from the recently warmed forest, probably due to insufficient evolutionary time, but springtails from the warmed grassland plots had significantly steeper reaction norms than their counterparts from the corresponding unwarmed plot. In contrast, cold tolerance and locomotory activity showed no conclusive clinal pattern despite significant within-habitat variation. There appeared to be significant differences between habitats, as springtails from the forest had more temperature-sensitive developmental rate and locomotory activity, walked faster, and exhibited more variable cold tolerance than grassland springtails did. The planting of a forest, therefore, seems to have exerted a stronger effect on the thermal phenotype of P. pseudovanderdrifti than the emergence of a geothermal gradient. Thus, habitat properties may be no less important in shaping thermal reaction norms than the mean temperature. These local-scale findings suggest that, in addition to warming per se, global transformation of communities may drive the evolution of thermal phenotypes to an extent comparable with the effect of rising environmental temperature

Phenological responses of Icelandic subarctic grasslands to short-term and long-term natural soil warming

The phenology of vegetation, particularly the length of the growing season (LOS; i.e., the period from greenup to senescence), is highly sensitive to climate change, which could imply potent feedbacks to the climate system, for example, by altering the ecosystem carbon (C) balance. In recent decades, the largest extensions of LOS have been reported at high northern latitudes, but further warming-induced LOS extensions may be constrained by too short photoperiod or unfulfilled chilling requirements. Here, we studied subarctic grasslands, which cover a vast area and contain large C stocks, but for which LOS changes under further warming are highly uncertain. We measured LOS extensions of Icelandic subarctic grasslands along natural geothermal soil warming gradients of different age (short term, where the measurements started after 5 years of warming and long term, i.e., warmed since ≥50 years) using ground-level measurements of normalized difference vegetation index. We found that LOS linearly extended with on average 2.1 days per °C soil warming up to the highest soil warming levels (ca. +10°C) and that LOS had the potential to extend at least 1 month. This indicates that the warming impact on LOS in these subarctic grasslands will likely not saturate in the near future. A similar response to short- and long-term warming indicated a strong physiological control of the phenological response of the subarctic grasslands to warming and suggested that genetic adaptations and community changes were likely of minor importance. We conclude that the warming-driven extension of the LOSs of these subarctic grasslands did not saturate up to +10°C warming, and hence that growing seasons of high-latitude grasslands are likely to continue lengthening with future warming (unless genetic adaptations or species shifts do occur). This persistence of the warming-induced extension of LOS has important implications for the C-sink potential of subarctic grasslands under climate change

Coupled carbon and nitrogen losses in response to seven years of chronic warming in subarctic soils

Increasing temperatures may alter the stoichiometric demands of soil microbes and impair their capacity to stabilize carbon (C) and retain nitrogen (N), with critical consequences for the soil C and N storage at high latitude soils. Geothermally active areas in Iceland provided wide, continuous and stable gradients of soil temperatures to test this hypothesis. In order to characterize the stoichiometric demands of microbes from these subarctic soils, we incubated soils from ambient temperatures after the factorial addition of C, N and P substrates separately and in combination. In a second experiment, soils that had been exposed to different in situ warming intensities (+0, +0.5, +1.8, +3.4, +8.7, +15.9 °C above ambient) for seven years were incubated after the combined addition of C, N and P to evaluate the capacity of soil microbes to store and immobilize C and N at the different warming scenarios. The seven years of chronic soil warming triggered large and proportional soil C and N losses (4.1 ± 0.5% °C−1 of the stocks in unwarmed soils) from the upper 10 cm of soil, with a predominant depletion of the physically accessible organic substrates that were weakly sorbed in soil minerals up to 8.7 °C warming. Soil microbes met the increasing respiratory demands under conditions of low C accessibility at the expenses of a reduction of the standing biomass in warmer soils. This together with the strict microbial C:N stoichiometric demands also constrained their capacity of N retention, and increased the vulnerability of soil to N losses. Our findings suggest a strong control of microbial physiology and C:N stoichiometric needs on the retention of soil N and on the resilience of soil C stocks from high-latitudes to warming, particularly during periods of vegetation dormancy and low C inputs

Impact of soil warming on the plant metabolome of Icelandic grasslands

Altres ajuts: Scholarly Studies programme of the Smithsonian Institution, projects LM2015061 and LO1415 of the Ministry of Education, Youth and Sports of the Czech Republic, and the Research Foundation-Flanders (FWO aspirant grant to N.L.).Climate change is stronger at high than at temperate and tropical latitudes. The natural geothermal conditions in southern Iceland provide an opportunity to study the impact of warming on plants, because of the geothermal bedrock channels that induce stable gradients of soil temperature. We studied two valleys, one where such gradients have been present for centuries (long-term treatment), and another where new gradients were created in 2008 after a shallow crustal earthquake (short-term treatment). We studied the impact of soil warming (0 to +15 °C) on the foliar metabolomes of two common plant species of high northern latitudes: Agrostis capillaris, a monocotyledon grass; and Ranunculus acris, a dicotyledonous herb, and evaluated the dependence of shifts in their metabolomes on the length of the warming treatment. The two species responded differently to warming, depending on the length of exposure. The grass metabolome clearly shifted at the site of long-term warming, but the herb metabolome did not. The main up-regulated compounds at the highest temperatures at the long-term site were saccharides and amino acids, both involved in heat-shock metabolic pathways. Moreover, some secondary metabolites, such as phenolic acids and terpenes, associated with a wide array of stresses, were also up-regulated. Most current climatic models predict an increase in annual average temperature between 2-8 °C over land masses in the Arctic towards the end of this century. The metabolomes of A. capillaris and R. acris shifted abruptly and nonlinearly to soil warming >5 °C above the control temperature for the coming decades. These results thus suggest that a slight warming increase may not imply substantial changes in plant function, but if the temperature rises more than 5 °C, warming may end up triggering metabolic pathways associated with heat stress in some plant species currently dominant in this region

Soil carbon loss in warmed subarctic grasslands is rapid and restricted to topsoil

Global warming may lead to carbon transfers from soils to the atmosphere, yet this positive feedback to the climate system remains highly uncertain, especially in subsoils (Ilyina and Friedlingstein, 2016; Shi et al., 2018). Using natural geothermal soil warming gradients of up to +6.4 degrees C in subarctic grasslands (Sigurdsson et al., 2016), we show that soil organic carbon (SOC) stocks decline strongly and linearly with warming (-2.8 t ha(-1) degrees C-1). Comparison of SOC stock changes following medium-term (5 and 10 years) and long-term (> 50 years) warming revealed that all SOC stock reduction occurred within the first 5 years of warming, after which continued warming no longer reduced SOC stocks. This rapid equilibration of SOC observed in Andosol suggests a critical role for ecosystem adaptations to warming and could imply short-lived soil carbon-climate feedbacks. Our data further revealed that the soil C loss occurred in all aggregate size fractions and that SOC stock reduction was only visible in topsoil (0-10 cm). SOC stocks in subsoil (10-30 cm), where plant roots were absent, showed apparent conservation after > 50 years of warming. The observed depth-dependent warming responses indicate that explicit vertical resolution is a prerequisite for global models to accurately project future SOC stocks for this soil type and should be investigated for soils with other mineralogies

Natural gradients in temperature and nitrogen: Iceland represents a unique environment to clarify long-term global change effects on carbon dynamics

Sameiginlegt með University of Antwerp - Faculty of ScienceLeblans, N. I. W. 2016. Natural gradients in temperature and nitrogen: Iceland represents a unique environment to clarify long-term global change effects on carbon dynamics. Joint doctoral thesis at Antwerp University and Agricultural University of Iceland. Global change is one of the greatest challenges of our generation. Surface temperatures are rising as a consequence of anthropogenic greenhouse gas (GHG) emissions, but projections remain highly uncertain as solid knowledge on potent feedback mechanisms from ecosystems to the climate system is limited. One such potentially powerful feedback mechanism is the warming-induced transfer of soil organic carbon (SOC) to the atmosphere, exacerbating the warming. This is especially true for high northern latitude ecosystems (≥ 60° N) where both the highest amount of SOC is stored and where warming is most pronounced. In this region, SOC storage is also strongly linked to nitrogen (N) cycling, as plant productivity, the primary source of SOC, is generally N limited in cold ecosystems. During the past decades, northern temperate and some boreal ecosystems have already been absorbing about 10% of the anthropogenic C emissions due to an N-driven increase in productivity. At high northern latitudes, the N inputs, which have remained relatively low up to now, are expected to increase substantially, potentially transforming northern ecosystems to important contributors of the N-induced C sink in the near future. However, the future evolution of both warminginduced SOC losses and the N-dependent C sink is highly uncertain due to the scarcity of empirical observations of long-term effects. In this thesis, natural gradients were used to obtain empirical observations of longterm warming and N input effects on C dynamics in subarctic grasslands, which cover ca. 10% of the global terrestrial surface. The primary advantage of natural gradients is that equilibrium states can be observed. Further, their typical wide and continuous nature allows to study the full range of global change projections and to detect non-linearities in the response. Here, I investigated natural (geothermal) soil temperature gradients (+0 - +20 °C) at the ForHot research site (www.forhot.is) in southwest Iceland, where the presence of both shortterm (formed in 2008) and long-term (≥ 50 years) gradients allowed to separate transient (short-term) from permanent (long-term) responses. Natural gradients in N inputs were investigated on islands of Vestmannaeyjar (off the south coast of Iceland) with differing soil age (and thus different N-accumulation time; 50 vs. 1,600 years) and amounts of seabird N inputs. Firstly, I aimed to evaluate the potential of natural gradients of geothermal soil warming and N inputs as global change laboratories and secondly to assess and compare their short-term and long-term effects on the C dynamics of subarctic grasslands, with focus on plant phenology and SOC sequestration. The geothermal soil temperature gradients proved to be a valuable complement to other warming techniques to improve our understanding of long-term warming effects on ecosystems, overcoming important difficulties that are typical for climate manipulation experiments, such as limited duration of the warming and a limited number of warming levels. The length of the growing season (LOS), a phenological parameter of great importance in the C balance of northern ecosystems, extended linearly with increasing soil temperature by on average 15.6 ± 4.7 (SE) days °C-1. This was primarily due to an advance in the start of the growing season, while the end of the growing season was largely unresponsive to warmer soil temperatures. This persistency of the warming-induced extension of LOS, which has recently been questioned, has important implications for the C-sink potential of subarctic grasslands under climate change. SOC stocks showed a large and linear decline under soil warming (4.1 ± 0.5 SE % °C- 1), both after 5 and ≥ 50 years of warming. Central to the observed SOC loss was a loss of physical stabilization in soil aggregates. This was an important finding and such mechanism is presently not embedded in any Earth System Model, but its inclusion could possibly improve model projections. Further, the similar SOC loss after short- and long-term warming suggested that warming of subarctic grassland soils could cause a rapid positive feedback to climate warming. In contrast to soil warming, increasing N inputs increased the short-term SOC storage rate of early successional and mature subarctic grasslands (from 0.018 to 0.29 and from 0.30 to 0.44 ton SOC ha-1 yr-1 respectively), as well as the long-term SOC storage rate of the mature grasslands (from 0.12 to 0.16 ton SOC ha-1 yr-1). This suggested that the N-induced C sink of subarctic grasslands could be maintained for many centuries, which contradicts the current hypothesis that the N-deposition driven terrestrial CO2 sink is likely to saturate in the near future

Phenological responses of Icelandic subarctic grasslands to short-term and long-term natural soil warming

The phenology of vegetation, particularly the length of the growing season (LOS; i.e., the period from greenup to senescence), is highly sensitive to climate change, which could imply potent feedbacks to the climate system, for example, by altering the ecosystem carbon (C) balance. In recent decades, the largest extensions of LOS have been reported at high northern latitudes, but further warming-induced LOS extensions may be constrained by too short photoperiod or unfulfilled chilling requirements. Here, we studied subarctic grasslands, which cover a vast area and contain large C stocks, but for which LOS changes under further warming are highly uncertain. We measured LOS extensions of Icelandic subarctic grasslands along natural geothermal soil warming gradients of different age (short term, where the measurements started after 5 years of warming and long term, i.e., warmed since ≥50 years) using ground-level measurements of normalized difference vegetation index. We found that LOS linearly extended with on average 2.1 days per °C soil warming up to the highest soil warming levels (ca. +10°C) and that LOS had the potential to extend at least 1 month. This indicates that the warming impact on LOS in these subarctic grasslands will likely not saturate in the near future. A similar response to short- and long-term warming indicated a strong physiological control of the phenological response of the subarctic grasslands to warming and suggested that genetic adaptations and community changes were likely of minor importance. We conclude that the warming-driven extension of the LOSs of these subarctic grasslands did not saturate up to +10°C warming, and hence that growing seasons of high-latitude grasslands are likely to continue lengthening with future warming (unless genetic adaptations or species shifts do occur). This persistence of the warming-induced extension of LOS has important implications for the C-sink potential of subarctic grasslands under climate change

Coupled carbon and nitrogen losses in response to seven years of chronic warming in subarctic soils

Increasing temperatures may alter the stoichiometric demands of soil microbes and impair their capacity to stabilize carbon (C) and retain nitrogen (N), with critical consequences for the soil C and N storage at high latitude soils. Geothermally active areas in Iceland provided wide, continuous and stable gradients of soil temperatures to test this hypothesis. In order to characterize the stoichiometric demands of microbes from these subarctic soils, we incubated soils from ambient temperatures after the factorial addition of C, N and P substrates separately and in combination. In a second experiment, soils that had been exposed to different in situ warming intensities (+0, +0.5, +1.8, +3.4, +8.7, +15.9 °C above ambient) for seven years were incubated after the combined addition of C, N and P to evaluate the capacity of soil microbes to store and immobilize C and N at the different warming scenarios. The seven years of chronic soil warming triggered large and proportional soil C and N losses (4.1 ± 0.5% °C−1 of the stocks in unwarmed soils) from the upper 10 cm of soil, with a predominant depletion of the physically accessible organic substrates that were weakly sorbed in soil minerals up to 8.7 °C warming. Soil microbes met the increasing respiratory demands under conditions of low C accessibility at the expenses of a reduction of the standing biomass in warmer soils. This together with the strict microbial C:N stoichiometric demands also constrained their capacity of N retention, and increased the vulnerability of soil to N losses. Our findings suggest a strong control of microbial physiology and C:N stoichiometric needs on the retention of soil N and on the resilience of soil C stocks from high-latitudes to warming, particularly during periods of vegetation dormancy and low C inputs

Impact of soil warming on the plant metabolome of Icelandic grasslands

Altres ajuts: Scholarly Studies programme of the Smithsonian Institution, projects LM2015061 and LO1415 of the Ministry of Education, Youth and Sports of the Czech Republic, and the Research Foundation-Flanders (FWO aspirant grant to N.L.).Climate change is stronger at high than at temperate and tropical latitudes. The natural geothermal conditions in southern Iceland provide an opportunity to study the impact of warming on plants, because of the geothermal bedrock channels that induce stable gradients of soil temperature. We studied two valleys, one where such gradients have been present for centuries (long-term treatment), and another where new gradients were created in 2008 after a shallow crustal earthquake (short-term treatment). We studied the impact of soil warming (0 to +15 °C) on the foliar metabolomes of two common plant species of high northern latitudes: Agrostis capillaris, a monocotyledon grass; and Ranunculus acris, a dicotyledonous herb, and evaluated the dependence of shifts in their metabolomes on the length of the warming treatment. The two species responded differently to warming, depending on the length of exposure. The grass metabolome clearly shifted at the site of long-term warming, but the herb metabolome did not. The main up-regulated compounds at the highest temperatures at the long-term site were saccharides and amino acids, both involved in heat-shock metabolic pathways. Moreover, some secondary metabolites, such as phenolic acids and terpenes, associated with a wide array of stresses, were also up-regulated. Most current climatic models predict an increase in annual average temperature between 2-8 °C over land masses in the Arctic towards the end of this century. The metabolomes of A. capillaris and R. acris shifted abruptly and nonlinearly to soil warming >5 °C above the control temperature for the coming decades. These results thus suggest that a slight warming increase may not imply substantial changes in plant function, but if the temperature rises more than 5 °C, warming may end up triggering metabolic pathways associated with heat stress in some plant species currently dominant in this region