Niki I. W. Leblans

Twenty-two years of warming, fertilisation and shading of subarctic heath shrubs promote secondary growth and plasticity but not primary growth.

Most manipulation experiments simulating global change in tundra were short-term or did not measure plant growth directly. Here, we assessed the growth of three shrubs (Cassiope tetragona, Empetrum hermaphroditum and Betula nana) at a subarctic heath in Abisko (Northern Sweden) after 22 years of warming (passive greenhouses), fertilisation (nutrients addition) and shading (hessian fabric), and compare this to observations from the first decade of treatment. We assessed the growth rate of current-year leaves and apical stem (primary growth) and cambial growth (secondary growth), and integrated growth rates with morphological measurements and species coverage. Primary- and total growth of Cassiope and Empetrum were unaffected by manipulations, whereas growth was substantially reduced under fertilisation and shading (but not warming) for Betula. Overall, shrub height and length tended to increase under fertilisation and warming, whereas branching increased mostly in shaded Cassiope. Morphological changes were coupled to increased secondary growth under fertilisation. The species coverage showed a remarkable increase in graminoids in fertilised plots. Shrub response to fertilisation was positive in the short-term but changed over time, likely because of an increased competition with graminoids. More erected postures and large, canopies (requiring enhanced secondary growth for stem reinforcement) likely compensated for the increased light competition in Empetrum and Cassiope but did not avoid growth reduction in the shade intolerant Betula. The impact of warming and shading on shrub growth was more conservative. The lack of growth enhancement under warming suggests the absence of long-term acclimation for processes limiting biomass production. The lack of negative effects of shading on Cassiope was linked to morphological changes increasing the photosynthetic surface. Overall, tundra shrubs showed developmental plasticity over the longer term. However, such plasticity was associated clearly with growth rate trends only in fertilised plots.

Stem Secondary Growth of Tundra Shrubs: Impact of Environmental Factors and Relationships with Apical Growth

Abstract Our knowledge of stem secondary growth of arctic shrubs (a key component of tundra net primary production, NPP) is very limited. Here, we investigated the impact of the physical elements of the environment on shrub secondary growth by comparing annual growth rates of model species from similar habitats at contrasting altitude, microtopography, latitude, geographical location, and soil type, in both the sub- and High Arctic. We found that secondary growth has a modest sensitivity to the environment but with large differences among species. For example, the evergreen Cassiope tetragona is affected by altitude, microtopography, and latitude, whereas the evergreen Empetrum hermaphroditum has rather constant secondary growth in all environments. Deciduous species seem to be most affected by microtopography. Furthermore, the impact of the environment on secondary growth differed from the impact on primary growth (stem apical growth, stem length, and apical growth of stem plus leaves), in some cases even with opposite responses. Thus caution should be taken when estimating the impact of the environment on shrub growth from apical growth only. Integration of our data set with the (very limited) previously published information on secondary growth provides an overview of its contribution to NPP and annual growth rates for 9 arctic species at 18 sites in Sweden, Greenland, Svalbard, Alaska, and the Alps.

Geothermal ecosystems as natural climate change experiments : The ForHot research site in Iceland as a case study

This article describes how natural geothermal soil temperature gradients in Iceland have been used to study terrestrial ecosystem responses to soil warming. The experimental approach was evaluated at three study sites in southern Iceland; one grassland site that has been warm for at least 50 years (GO), and another comparable grassland site (GN) and a Sitka spruce plantation (FN) site that have both been warmed since an earthquake took place in 2008. Within each site type, five ca. 50 m long transects, with six permanent study plots each, were established across the soil warming gradients, spanning from unwarmed control conditions to gradually warmer soils. It was attempted to select the plots so the annual warming levels would be ca. +1, +3, +5, +10 and +20 °C within each transect. Results of continuous measurements of soil temperature (Ts) from 2013-2015 revealed that the soil warming was relatively constant and followed the seasonal Ts cycle of the unwarmed control plots. Volumetric water content in the top 5 cm of soil was repeatedly surveyed during 2013-2016. The grassland soils were wetter than the FN soils, but they had sometimes some significant warming-induced drying in the surface layer of the warmest plots, in contrast to FN. Soil chemistry did not show any indications that geothermal water had reached the root zone, but soil pH did increase somewhat with warming, which was probably linked to vegetation changes. As expected, the potential decomposition rate of organic matter increased significantly with warming. It was concluded that the natural geothermal gradients at the ForHot sites in Iceland offered realistic conditions for studying terrestrial ecosystem responses to warming with minimal artefacts.

Prolonged exposure does not increase soil microbial community compositional response to warming along geothermal gradients

ABSTRACT Global change is expected to affect soil microbial communities through their responsiveness to temperature. It has been proposed that prolonged exposure to elevated temperatures may lead to progressively larger effects on soil microbial community composition. However, due to the relatively short‐term nature of most warming experiments, this idea has been challenging to evaluate. The present study took the advantage of natural geothermal gradients (from +1°C to +19°C above ambient) in two subarctic grasslands to test the hypothesis that long‐term exposure (>50 years) intensifies the effect of warming on microbial community composition compared to short‐term exposure (5‐7 years). Community profiles from amplicon sequencing of bacterial and fungal rRNA genes did not support this hypothesis: significant changes relative to ambient were observed only starting from the warming intensity of +9°C in the long term and +7°C/+3°C in the short term, for bacteria and fungi, respectively. Our results suggest that microbial communities in high‐latitude grasslands will not undergo lasting shifts in community composition under the warming predicted for the coming 100 years (+2.2°C to +8.3°C).

Microbial temperature sensitivity and biomass change explain soil carbon loss with warming

Soil microorganisms control carbon losses from soils to the atmosphere1–3, yet their responses to climate warming are often short-lived and unpredictable4–7. Two mechanisms, microbial acclimation and substrate depletion, have been proposed to explain temporary warming effects on soil microbial activity8–10. However, empirical support for either mechanism is unconvincing. Here we used geothermal temperature gradients (>50 years of field warming)11 and a short-term experiment to show that microbial activity (gross rates of growth, turnover, respiration and carbon uptake) is intrinsically temperature sensitive and does not acclimate to warming (+6 °C) over weeks or decades. Permanently accelerated microbial activity caused carbon loss from soil. However, soil carbon loss was temporary because substrate depletion reduced microbial biomass and constrained the influence of microbes over the ecosystem. A microbial biogeochemical model12–14 showed that these observations are reproducible through a modest, but permanent, acceleration in microbial physiology. These findings reveal a mechanism by which intrinsic microbial temperature sensitivity and substrate depletion together dictate warming effects on soil carbon loss via their control over microbial biomass. We thus provide a framework for interpreting the links between temperature, microbial activity and soil carbon loss on timescales relevant to Earth’s climate system.Soil microbial activity is accelerated by warming and does not acclimate over periods of at least 50 years. Resulting soil carbon loss is nevertheless temporary because substrate depletion reduces microbial biomass and constrains the influence of microbes over the ecosystem.

Impact of Soil Warming on the Plant Metabolome of Icelandic Grasslands

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.

Author Correction: Microbial temperature sensitivity and biomass change explain soil carbon loss with warming

In the version of this Letter originally published, the name of the institute in affiliation 3 was incorrect; it read “Institute of Applied Systems Analysis” but should have read “International Institute for Applied Systems Analysis”. This has now been corrected.

Functional diversity of Collembola is reduced in soils subjected to short‐term, but not long‐term, geothermal warming

1. Human activities have caused global changes of atmospheric chemistry resulting in increased temperature especially in the colder regions of the northern hemisphere. Since warming of the environment can have drastic effects on terrestrial ecosystems it is important to experimentally evaluate the extent of such effects in long-term field-based experiments. In this study we make use of both recent (short-term) and long-term geothermal warming of Icelandic soils to examine the responses of Collembola, an ecologically important group of soil invertebrates, to warming. 2. On the basis of metabolic scaling theory, we hypothesized that species of small size would be more successful in warmed soils than species of larger size. Further we expected that top-soil-dwelling species would benefit more from warming than deep-soil-dwelling species. In order to test these hypotheses we sampled Collembola along replicated gradients of increasing temperature in areas that had been heated for about 6 years and more than 50 years respectively. Collembola were identified to species level, counted and the community-weighted mean trait scores for six functional and ecological traits were calculated. 3. Results show that both short-term and long-term soil warming caused a shift towards a higher relative abundance of species with small body size. Furthermore, abundance of top-soil-dwelling Collembola tended to increase after short-term warming, but the opposite was observed after long-term warming. 4. Using trait-based diversity indices (FRic and RaoQ), we show that functional richness and diversity of Collembola communities was significantly reduced (almost halved) as a result of short-term soil warming to about 10 degrees C above normal, but this effect was not detected in plots equally warmed for more than 50 years. This indicates that the functional diversity of Collembola communities have high resilience towards soil warming in a long-term perspective.

Acclimation of Fine Root Systems to Soil Warming: Comparison of an Experimental Setup and a Natural Soil Temperature Gradient

Global warming is predicted to impact high-latitude and high-altitude forests severely, jeopardizing their overall functioning and carbon storage, both of which depend on the warming response of tree fine root systems. This paper investigates the effect of soil warming on the biomass, morphology and colonizing ectomycorrhizal community of spruce fine and absorptive fine roots. We compare the responses of spruce roots growing at a man-made long-term soil warming (+ 4°C) experiment to results obtained from a geothermal soil temperature gradient (+ 1 to + 14°C) extending to the forest die-off edge, to shed light on the generalizability of the warming response and reveal any thresholds in acclimation ability. Trees in warmer soils formed longer and less-branched absorptive roots with higher specific root length and area, and lower root tissue density in both spruce stands, irrespective of warming method and location. Soil warming at the experimental warming site also supported the occurrence of a more varied EcM community and an increase in the abundance of Tomentella spp., indicating a shift in nutrient foraging. Fine and absorptive fine root biomass decreased toward warmer soil, with a sharp reduction occurring between + 4 and + 6°C from the ambient and leading to the collapse of the fine root system at the geothermal gradient. At the experimental warming site, the applied + 4°C warming had no effect on fine and absorptive fine root biomass. The similar fine root responses at the two warming sites suggest that the observations possibly reflect general acclimation patterns in spruce forests to global warming.