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  • 1. Balogianni, Vasiliki G.
    et al.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Wilson, Scott D.
    Root production in contrasting ecosystems: the impact of rhizotron sampling frequency2016In: Plant Ecology, ISSN 1385-0237, E-ISSN 1573-5052, Vol. 217, no 11, p. 1359-1367Article in journal (Refereed)
    Abstract [en]

    Despite their critical role in every terrestrial ecosystem, fine root production and mortality have not been widely compared among systems due to the practical difficulties of belowground research. We examined fine root production and mortality among five contrasting sites: native and invaded grassland in eastern Montana, USA, aspen forest in southern Saskatchewan, Canada, and birch forest and tundra in northern Sweden. Additionally, we investigated the importance of minirhizotron sampling interval on measures of root production and mortality by comparing measures produced from 1-, 7-, 14-, and 21-day sample intervals. Root length and mortality varied significantly among sites, with invaded grassland having the greatest root length (> 2 x than any other site) and significantly greater root mortality than native grassland (54 %). In contrast, there were no significant differences in root production among the sites. Sample interval had no significant influence on root production or mortality. Minirhizotron sampling intervals up to 3 weeks did not underestimate the measures of root production and mortality in comparison to measures derived from shorter sampling intervals, regardless of the site studied. The results suggest that 3 weeks can be an accurate and efficient sample interval when studying root production and mortality with minirhizotrons.

  • 2.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    The belowground growing season2022In: Nature Climate Change, ISSN 1758-678X, E-ISSN 1758-6798, Vol. 12, p. 11-12Article in journal (Other academic)
  • 3.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    The hidden life of plants: fine root dynamics in northern ecosystems2016Doctoral thesis, comprehensive summary (Other academic)
    Abstract [en]

    Fine roots constitute a large part of the primary production in northern (arctic and boreal) ecosystems, and are key players in ecosystem fluxes of water, nutrients and carbon. Data on root dynamics are generally rare, especially so in northern ecosystems. However, those ecosystems undergo the most rapid climatic changes on the planet and a profound understanding of form, function and dynamics of roots in such ecosystems is essential.

    This thesis aimed to advance our knowledge about fine root dynamics in northern ecosystems, with a focus on fine root phenology in natural plant communities and how climate change might alter it. Factors considered included thickness and duration of snow cover, thawing of permafrost, as well as natural gradients in temperature. Experiments and observational studies were located around Abisko (68°21' N, 18°45' E), and in a boreal forest close to Vindeln (64°14'N, 19°46'E), northern Sweden. Root responses included root growth, total root length, and root litter input, always involving seasonal changes therein, measured with minirhizotrons. Root biomass was also determined with destructive soil sampling. Additionally, aboveground response parameters, such as phenology and growth, and environmental parameters, such as air and soil temperatures, were assessed.

    This thesis reveals that aboveground patterns or responses cannot be directly translated belowground and urges a decoupling of above- and belowground phenology in terrestrial biosphere models. Specifically, root growth occurred outside of the photosynthetically active period of tundra plants. Moreover, patterns observed in arctic and boreal ecosystems diverged from those of temperate systems, and models including root parameters may thus need specific parameterization for northern ecosystems. In addition, this thesis showed that plant communities differ in root properties, and that changes in plant community compositions can thus induce changes in root dynamics and functioning. This underlines the importance of a thorough understanding of root dynamics in different plant community types in order to understand and predict how changes in plant communities in response to climate change will translate into root dynamics. Overall, this thesis describes root dynamics in response to a variety of factors, because a deeper knowledge about root dynamics will enable a better understanding of ecosystem processes, as well as improve model prediction of how northern ecosystems will respond to climate change.

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  • 4.
    Blume-Werry, Gesche
    et al.
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Di Maurizio, Vanessa
    Beil, Ilka
    Lett, Signe
    Schwieger, Sarah
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Kreyling, Juergen
    Don't drink it, bury it: comparing decomposition rates with the tea bag index is possible without prior leaching2021In: Plant and Soil, ISSN 0032-079X, E-ISSN 1573-5036, Vol. 465, no 1-2, p. 613-621Article in journal (Refereed)
    Abstract [en]

    Purpose: The standardized ‘Tea Bag Index’ enables comparisons of litter decomposition rates, a key component of carbon cycling, across ecosystems. However, tea ‘litter’ may leach more than other plant litter, skewing comparisons of decomposition rates between sites with differing moisture conditions. Therefore, some researchers leach tea bags before field incubation. This decreases comparability between studies, and it is unclear if this modification is necessary.

    Methods: We submerged green and rooibos tea bags in water, and measured their leaching losses over time (2 min – 72 h). We also compared leaching of tea to leaf and root litter from other plant species, and finally, compared mass loss of pre-leached and standard tea bags in a fully factorial incubation experiment differing in soil moisture (wet and dry) and soil types (sand and peat).

    Results: Both green and rooibos tea leached strongly, levelling-off at about 40% and 20% mass loss, respectively. Mass loss from leaching was highest in green tea followed by leaves of other plants, then rooibos tea, and finally roots of other plants. When incubated for 4 weeks, both teas showed lower mass loss when they had been pre-leached compared to standard tea bags. However, these differences between standard and pre-leached tea bags were similar in moist vs. dry soils, both in peat and in sand.

    Conclusions: Thus, despite large leaching losses, we conclude that leaching tea bags before field or lab incubation is not necessary to compare decomposition rates between systems, ranging from as much as 5% to 25% soil moisture.

  • 5.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Dorrepaal, Ellen
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Keuper, Frida
    BioEcoAgro Joint Research Unit, INRAE, Barenton-Bugny, France.
    Kummu, Matti
    Water and Development Research Group, Aalto University, Aalto, Finland.
    Wild, Birgit
    Department of Environmental Science, Stockholm University, Stockholm, Sweden; Bolin Centre for Climate Research, Stockholm University, Stockholm, Sweden.
    Weedon, James T.
    Amsterdam Institute for Life and Environment (A-LIFE), Systems Ecology Section, Vrije Universiteit Amsterdam, Amsterdam, Netherlands.
    Arctic rooting depth distribution influences modelled carbon emissions but cannot be inferred from aboveground vegetation type2023In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 240, no 2, p. 502-514Article in journal (Refereed)
    Abstract [en]

    The distribution of roots throughout the soil drives depth-dependent plant–soil interactions and ecosystem processes, particularly in arctic tundra where plant biomass, is predominantly belowground. Vegetation is usually classified from aboveground, but it is unclear whether such classifications are suitable to estimate belowground attributes and their consequences, such as rooting depth distribution and its influence on carbon cycling. We performed a meta-analysis of 55 published arctic rooting depth profiles, testing for differences both between distributions based on aboveground vegetation types (Graminoid, Wetland, Erect-shrub, and Prostrate-shrub tundra) and between ‘Root Profile Types’ for which we defined three representative and contrasting clusters. We further analyzed potential impacts of these different rooting depth distributions on rhizosphere priming-induced carbon losses from tundra soils. Rooting depth distribution hardly differed between aboveground vegetation types but varied between Root Profile Types. Accordingly, modelled priming-induced carbon emissions were similar between aboveground vegetation types when they were applied to the entire tundra, but ranged from 7.2 to 17.6 Pg C cumulative emissions until 2100 between individual Root Profile Types. Variations in rooting depth distribution are important for the circumpolar tundra carbon-climate feedback but can currently not be inferred adequately from aboveground vegetation type classifications.

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  • 6.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Jansson, Roland
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Root phenology unresponsive to earlier snowmelt despite advanced aboveground phenology in two subarctic plant communitiesManuscript (preprint) (Other academic)
    Abstract [en]

    Earlier snowmelt at high latitudes advances aboveground plant phenology, thereby affecting water, nutrient and carbon cycles. Despite the key role of fine roots in these ecosystem processes, phenological responses to earlier snowmelt have never been assessed belowground. We experimentally advanced snowmelt in two contrasting plant community types (heath and meadow) in northern Sweden and measured above- and belowground phenology (leaf-out, flowering and fine root growth). We expected earlier snowmelt to advance both above- and belowground phenology, and shrub-dominated heath to be more responsive than meadow. Snow melted on average nine days earlier in the manipulated plots than in controls, and soil temperatures were on average 0.9 °C higher during the snowmelt period of three weeks. This resulted in small advances in aboveground phenology, but contrary to our expectations, root phenology was unresponsive, with root growth generally starting before leaf-out. Both plant community types responded similarly to the snowmelt treatment, despite strong differences in dominating plant functional types, and root properties, such as root length and turnover. The lack of a response in root phenology, despite warmer soil temperatures and aboveground phenological advances, adds evidence that aboveground plant responses might not be directly translated to belowground plant responses, and that our understanding of factors driving belowground phenology is still limited, although of major importance for water, nutrient and carbon cycling.

  • 7.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Jansson, Roland
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Biodiversity and Natural Environment, Research Institute for Nature and Forest INBO, Kliniekstraat 25,1070 Brussels, Belgium.
    Root phenology unresponsive to earlier snowmelt despite advanced above-ground phenology in two subarctic plant communities2017In: Functional Ecology, ISSN 0269-8463, E-ISSN 1365-2435, Vol. 31, no 7, p. 1493-1502Article in journal (Refereed)
    Abstract [en]

    1. Earlier snowmelt at high latitudes advances above-ground plant phenology, thereby affecting water, nutrient and carbon cycles. Despite the key role of fine roots in these ecosystem processes, phenological responses to earlier snowmelt have never been assessed below-ground. 2. We experimentally advanced snowmelt in two contrasting plant community types (heath and meadow) in northern Sweden and measured above- and below-ground phenology (leaf-out, flowering and fine root growth). We expected earlier snowmelt to advance both above- and below-ground phenology, and shrub-dominated heath to be more responsive than meadow. 3. Snow melted on average 9 days earlier in the manipulated plots than in controls, and soil temperatures were on average 0.9 degrees C higher during the snowmelt period of 3 weeks. This resulted in small advances in above-ground phenology, but contrary to our expectations, root phenology was unresponsive, with root growth generally starting before leaf-out. These responses to the snowmelt treatment were similar in both plant community types, despite strong differences in dominating plant functional types and root properties, such as root length and turnover. 4. The lack of a response in root phenology, despite warmer soil temperatures and above-ground phenological advances, adds evidence that above-ground plant responses might not be directly translated to below-ground plant responses, and that our understanding of factors driving below-ground phenology is still limited, although of major importance for water, nutrient and carbon cycling.

  • 8.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Klaminder, Jonatan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Krab, Eveline J
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Soil and Environment, Swedish University of Agricultural Sciences, Uppsala, Sweden.
    Onteux, Sylvain
    Department of Environmental Science, Stockholm University, Stockholm, Sweden; Bolin Center for Climate Research, Stockholm University, Stockholm, Sweden.
    Ideas and perspectives: Alleviation of functional limitations by soil organisms is key to climate feedbacks from arctic soils2023In: Biogeosciences, ISSN 1726-4170, E-ISSN 1726-4189, Vol. 20, no 10, p. 1979-1990Article in journal (Refereed)
    Abstract [en]

    Arctic soils play an important role in Earth's climate system, as they store large amounts of carbon that, if released, could strongly increase greenhouse gas levels in our atmosphere. Most research to date has focused on how the turnover of organic matter in these soils is regulated by abiotic factors, and few studies have considered the potential role of biotic regulation. However, arctic soils are currently missing important groups of soil organisms, and here, we highlight recent empirical evidence that soil organisms' presence or absence is key to understanding and predicting future climate feedbacks from arctic soils. We propose that the arrival of soil organisms into arctic soils may introduce "novel functions", resulting in increased rates of, for example, nitrification, methanogenesis, litter fragmentation, or bioturbation, and thereby alleviate functional limitations of the current community. This alleviation can greatly enhance decomposition rates, in parity with effects predicted due to increasing temperatures. We base this argument on a series of emerging experimental evidence suggesting that the dispersal of until-then absent micro-, meso-, and macroorganisms (i.e. from bacteria to earthworms) into new regions and newly thawed soil layers can drastically affect soil functioning. These new observations make us question the current view that neglects organism-driven "alleviation effects"when predicting future feedbacks between arctic ecosystems and our planet's climate. We therefore advocate for an updated framework in which soil biota and the functions by which they influence ecosystem processes become essential when predicting the fate of soil functions in warming arctic ecosystems.

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  • 9.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, Soldmannstraße, Greifswald, Germany.
    Krab, Eveline J
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Swedish University of Agricultural Sciences, Department of Soil and Environment, Uppsala, Sweden.
    Olofsson, Johan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Sundqvist, Maja K.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Earth Sciences, University of Gothenburg, Gothenburg, Sweden.
    Väisänen, Maria
    Klaminder, Jonatan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Invasive earthworms unlock arctic plant nitrogen limitation2020In: Nature Communications, E-ISSN 2041-1723, Vol. 11, no 1Article in journal (Refereed)
    Abstract [en]

    Arctic plant growth is predominantly nitrogen (N) limited. This limitation is generally attributed to slow soil microbial processes due to low temperatures. Here, we show that arctic plant-soil N cycling is also substantially constrained by the lack of larger detritivores (earthworms) able to mineralize and physically translocate litter and soil organic matter. These new functions provided by earthworms increased shrub and grass N concentration in our common garden experiment. Earthworm activity also increased either the height or number of floral shoots, while enhancing fine root production and vegetation greenness in heath and meadow communities to a level that exceeded the inherent differences between these two common arctic plant communities. Moreover, these worming effects on plant N and greening exceeded reported effects of warming, herbivory and nutrient addition, suggesting that human spreading of earthworms may lead to substantial changes in the structure and function of arctic ecosystems. Arctic plant growth is predominantly nitrogen limited, where the slow nitrogen turnover in the soil is commonly attributed to the cold arctic climate. Here the authors show that the arctic plant-soil nitrogen cycling is also constrained by the lack of larger detritivores like earthworms.

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  • 10.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Kreyling, Juergen
    Greifswald University.
    Laudon, Hjalmar
    Sveriges Lantbruksuniversitet Umeå.
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Research Institute for Nature and Forest INBO, Kliniekstraat 25, 1070 Brussels, Belgium.
    Short-term climate change manipulation effects do not scale up to long-term legacies: effects of an absent snow cover on boreal forest plants2016In: Journal of Ecology, ISSN 0022-0477, E-ISSN 1365-2745, Vol. 104, no 6, p. 1638-1648Article in journal (Refereed)
    Abstract [en]

    1. Despite time-lags and nonlinearity in ecological processes, the majority of our knowledge about ecosystem responses to long-term changes in climate originates from relatively short-term experiments.

    2. We utilized the longest ongoing snow removal experiment in the world and an additional set of new plots at the same location in northern Sweden to simultaneously measure the effects of longterm (11 winters) and short-term (1 winter) absence of snow cover on boreal forest understorey plants, including the effects on root growth and phenology.

    3. Short-term absence of snow reduced vascular plant cover in the understorey by 42%, reduced fine root biomass by 16%, reduced shoot growth by up to 53% and induced tissue damage on two common dwarf shrubs. In the long-term manipulation, more substantial effects on understorey plant cover (92% reduced) and standing fine root biomass (39% reduced) were observed, whereas other response parameters, such as tissue damage, were observed less. Fine root growth was generally reduced, and its initiation delayed by c. 3 (short-term) to 6 weeks (long-term manipulation).

    4. Synthesis. We show that one extreme winter with a reduced snow cover can already induce ecologically significant alterations. We also show that long-term changes were smaller than suggested by an extrapolation of short-term manipulation results (using a constant proportional decline). In addition, some of those negative responses, such as frost damage and shoot growth, were even absolutely stronger in the short-term compared to the long-term manipulation. This suggests adaptation or survival of only those individuals that are able to cope with these extreme winter conditions, and that the short-term manipulation alone would overpredict long-term impacts. These results highlight both the ecological importance of snow cover in this boreal forest, and the value of combining short- and long-term experiments side by side in climate change research.

  • 11.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Lindén, Elin
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Andresen, Lisa
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Classen, Aimée T.
    Sanders, Nathan J.
    von Oppen, Jonathan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Sundqvist, Maja K.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Center for Macroecology, Evolution and Climate, The Natural History Museum of Denmark, University of Copenhagen, Copenhagen K, Denmark.
    Proportion of fine roots, but not plant biomass allocation below ground, increases with elevation in arctic tundra2018In: Journal of Vegetation Science, ISSN 1100-9233, E-ISSN 1654-1103, Vol. 29, no 2, p. 226-235Article in journal (Refereed)
    Abstract [en]

    Questions: Roots represent a considerable proportion of biomass, primary production and litter input in arctic tundra, and plant allocation of biomass to above- or below-ground tissue in response to climate change is a key factor in the future C balance of these ecosystems. According to optimality theory plants allocate C to the above- or below-ground structure that captures the most limiting resource. We used an elevational gradient to test this theory and as a space-for-time substitution to inform on tundra carbon allocation patterns under a shifting climate, by exploring if increasing elevation was positively related to the root:shoot ratio, as well as a larger plant allocation to adsorptive over storage roots.

    Location: Arctic tundra heath dominated by Empetrum hermaphroditum close to Abisko, Sweden.

    Methods: We measured root:shoot and fine:coarse root ratios of the plant communities along an elevational gradient by sampling above- and below-ground biomass, further separating root biomass into fine (<1 mm) and coarse roots.

    Results: Plant biomass was higher at the lower elevations, but the root:shoot ratio did not vary with elevation. Resource allocation to fine relative to coarse roots increased with elevation, resulting in a fine:coarse root ratio that more than doubled with increasing elevation.

    Conclusions: Contrary to previous works, the root:shoot ratio along this elevational gradient remained stable. However, communities along our study system were dominated by the same species at each elevation, which suggests that when changes in the root:shoot ratio occur with elevation these changes may be driven by differences in allocation patterns among species and thus turnover in plant community structure. Our results further reveal that the allocation of biomass to fine relative to coarse roots can differ between locations along an elevational gradient, even when overall above- vs below-ground biomass allocation does not. Given the functionally different roles of fine vs coarse roots this could have large implications for below-ground C cycling. Our results highlight the importance of direct effects vs indirect effects (such as changes in plant community composition and nutrient availability) of climate change for future C allocation above and below ground.

  • 12.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Germany.
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Research Institute for Nature and Forest INBO, Brussels, Belgium.
    Teuber, Laurenz M.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald, Germany.
    Johansson, Margareta
    Dorrepaal, Ellen
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Dwelling in the deep – strongly increased root growth and rooting depth enhance plant interactions with thawing permafrost soil2019In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 223, no 3, p. 1328-1339Article in journal (Refereed)
    Abstract [en]

    Climate‐warming‐induced permafrost thaw exposes large amounts of carbon and nitrogen in soil at considerable depths, below the seasonally thawing active layer. The extent to which plant roots can reach and interact with these hitherto detached, deep carbon and nitrogen stores remains unknown.

    We aimed to quantify how permafrost thaw affects root dynamics across soil depths and plant functional types compared with above‐ground abundance, and potential consequences for plant–soil interactions.

    A decade of experimental permafrost thaw strongly increased total root length and growth in the active layer, and deep roots invaded the newly thawed permafrost underneath. Root litter input to soil across all depths was 10 times greater with permafrost thaw. Root growth timing was unaffected by experimental permafrost thaw but peaked later in deeper soil, reflecting the seasonally receding thaw front. Deep‐rooting species could sequester 15N added at the base of the ambient active layer in October, which was after root growth had ceased.

    Deep soil organic matter that has long been locked up in permafrost is thus no longer detached from plant processes upon thaw. Whether via nutrient uptake, carbon storage, or rhizosphere priming, plant root interactions with thawing permafrost soils may feed back on our climate both positively and negatively.

  • 13.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Teuber, Laurenz M.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Margareta, Johansson
    Lund University.
    Dorrepaal, Ellen
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Dwelling in the deep – permafrost thawing strongly increases plant root growth and root litter inputManuscript (preprint) (Other academic)
    Abstract [en]

    Plant roots play a key role in ecosystem carbon and nutrient cycling. Climate warming induced thawing of permafrost exposes large amounts of carbon and nitrogen at greater soil depths that hitherto have been detached from plant influences. Whether plant roots can reach and interact with these carbon and nitrogen sources upon permafrost thaw remains unknown. Here, we use a long-term permafrost thaw experiment and a short-term deep fertilization experiment in northern Sweden to assess changes in vegetation composition and root dynamics (deep nitrogen uptake, root depth distribution, root growth and phenology, root mortality and litter input) related to permafrost thaw, both in active layer and in newly thawed permafrost. We show that Eriophorum vaginatum and Rubus chamaemorus, both relatively deep-rooting species, can take up nitrogen released at depth of permafrost thaw, despite the late release time in autumn when plant activity is expected to have ceased. Also, root dynamics changed drastically after a decade of experimental permafrost thaw. Total root length, root growth and root litter input all strongly increased, not only in the active layer but also in the newly thawed permafrost, and the timing of root growth was related to the seasonality of soil thaw. These responses were driven by Eriophorum vaginatum, which differed greatly in root dynamics compared to the other species and thus worked as an ecosystem engineer. This study demonstrates that soil organic matter currently locked-up at depth in permafrost is no longer detached from plant processes upon thaw. Given the pivotal role that roots have in the carbon cycle and the importance of the large carbon stocks in arctic soils, the changes observed here have the potential to feedback onto the global climate system.

  • 14.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Semenchuk, Philipp
    Department of Arctic Biology, UNIS – The University Centre in Svalbard, Longyearbyen, Norway.
    Ljung, Karin
    Department of Forest Genetics and Plant Physiology, Umeå Plant Science Centre, Swedish University of Agricultural Sciences, Umeå, Sweden.
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Novak, Ondrej
    Department of Forest Genetics and Plant Physiology, Umeå Plant Science Centre, Swedish University of Agricultural Sciences, Umeå, Sweden; Laboratory of Growth Regulators, Faculty of Science, Palacký University & Institute of Experimental Botany of the Czech Academy of Sciences, Olomouc, Czech Republic.
    Olofsson, Johan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Brunoni, Federica
    Department of Forest Genetics and Plant Physiology, Umeå Plant Science Centre, Swedish University of Agricultural Sciences, Umeå, Sweden; Laboratory of Growth Regulators, Faculty of Science, Palacký University & Institute of Experimental Botany of the Czech Academy of Sciences, Olomouc, Czech Republic.
    In situ seasonal patterns of root auxin concentrations and meristem length in an arctic sedge2024In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137Article in journal (Refereed)
    Abstract [en]
    • Seasonal dynamics of root growth play an important role in large-scale ecosystem processes; they are largely governed by growth regulatory compounds and influenced by environmental conditions. Yet, our knowledge about physiological drivers of root growth is mostly limited to laboratory-based studies on model plant species.
    • We sampled root tips of Eriophorum vaginatum and analyzed their auxin concentrations and meristem lengths biweekly over a growing season in situ in a subarctic peatland, both in surface soil and at the permafrost thawfront.
    • Auxin concentrations were almost five times higher in surface than in thawfront soils and increased over the season, especially at the thawfront. Surprisingly, meristem length showed an opposite pattern and was almost double in thawfront compared with surface soils. Meristem length increased from peak to late season in the surface soils but decreased at the thawfront.
    • Our study of in situ seasonal dynamics in root physiological parameters illustrates the potential for physiological methods to be applied in ecological studies and emphasizes the importance of in situ measurements. The strong effect of root location and the unexpected opposite patterns of meristem length and auxin concentrations likely show that auxin actively governs root growth to ensure a high potential for nutrient uptake at the thawfront.
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  • 15.
    Blume-Werry, Gesche
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Wilson, Scott D.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Biology, University of Regina, Regina, Saskatchewan S4S 0A2 Canada.
    Kreyling, Juergen
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Research Institute for Nature and Forest INBO, Kliniekstraat 25, 1070 Brussels, Belgium.
    The hidden season: growing season is 50% longer below than above ground along an arctic elevation gradient2016In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 209, no 3, p. 978-986Article in journal (Refereed)
    Abstract [en]

    There is compelling evidence from experiments and observations that climate warming prolongs the growing season in arctic regions. Until now, the start, peak, and end of the growing season, which are used to model influences of vegetation on biogeochemical cycles, were commonly quantified using above-ground phenological data. Yet, over 80% of the plant biomass in arctic regions can be below ground, and the timing of root growth affects biogeochemical processes by influencing plant water and nutrient uptake, soil carbon input and microbial activity. We measured timing of above- and below-ground production in three plant communities along an arctic elevation gradient over two growing seasons. Below-ground production peaked later in the season and was more temporally uniform than above-ground production. Most importantly, the growing season continued c. 50% longer below than above ground. Our results strongly suggest that traditional above-ground estimates of phenology in arctic regions, including remotely sensed information, are not as complete a representation of whole-plant production intensity or duration, as studies that include root phenology. We therefore argue for explicit consideration of root phenology in studies of carbon and nutrient cycling, in terrestrial biosphere models, and scenarios of how arctic ecosystems will respond to climate warming.

  • 16. De Long, Jonathan R.
    et al.
    Laudon, Hjalmar
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Kardol, Paul
    Nematode community resistant to deep soil frost in boreal forest soils2016In: Pedobiologia, ISSN 0031-4056, E-ISSN 1873-1511, Vol. 59, no 5-6, p. 243-251Article in journal (Refereed)
    Abstract [en]

    As global climate change advances, shifts in winter precipitation are becoming more common in high latitude ecosystems, resulting in less insulating snow cover and deeper soil frost. Long-term alterations to soil frost can impact on ecosystem processes such as decomposition, microbial activity and vegetation dynamics. In this study we utilized the longest running, well-characterized soil frost manipulation experiment in a boreal forest. We measured nematode family composition and feeding group abundances at four different soil layer depths from plots that had been subjected to deep soil frost for one and 11 years. The overall abundance of nematodes and the different feeding groups were unaffected by deep soil frost. However, a higher Maturity Index was weakly associated with deep soil frost (indicative of lower nutrient enrichment and more persister nematode (i.e., K-strategist) families), likely due to the loss of nutrients and reduced inputs from inhibited decomposition. Multivariate and regression analyses showed that most nematode families were weakly associated with dominant understory plant species and strongly associated with soil organic matter (SOM). This is probably the result of higher resource availability in the control plots, which is favorable to the nematode community. These results indicate that the nematode community was more strongly driven by the long-term indirect effects of deep soil frost on SOM as opposed to the direct effects. Our findings highlight that the indirect effects of altered winter precipitation and soil frost patterns may be more important than direct winter climate effects. Further, such indirect effects on SOM and the plant community that may affect the nematode community can only be seen in long-term experiments. Finally, given the critical role nematodes play in soil food webs and carbon and nutrient cycling, our results demonstrate the necessity of considering the response of nematodes to global climate change in boreal forest soils. 

  • 17.
    Gillert, Alexander
    et al.
    Fraunhofer Institute for Computer Graphics Research Igd, Rostock, Germany.
    Peters, Bo
    Greifswald University, Institute of Botany and Landscape Ecology, Germany.
    Von Lukas, Uwe Freiherr
    Fraunhofer Institute for Computer Graphics Research Igd, Rostock, Germany; University of Rostock, Institute for Visual... Analytic Computing, Germany.
    Kreyling, Jurgen
    Fraunhofer Institute for Computer Graphics Research Igd, Rostock, Germany.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Greifswald University, Institute of Botany and Landscape Ecology, Germany.
    Tracking growth and decay of plant roots in minirhizotron images2023In: Proceedings - 2023 IEEE Winter Conference on Applications of Computer Vision, WACV 2023, Institute of Electrical and Electronics Engineers (IEEE), 2023, p. 3688-3697Conference paper (Refereed)
    Abstract [en]

    Plant roots are difficult to monitor and study since they are hidden belowground. Minirhizotrons offer an in-situ monitoring solution but their widespread adoption is still limited by the capabilities of automatic analysis methods. These capabilities so far consist only of estimating a single number (total root length) per image.We propose a method for a more fine-grained analysis which estimates the root turnover, i.e. the amount of root growth and decay between two minirhizotron images. It consists of a neural network that computes which roots are visible in both images and is trained in an unsupervised manner without additional annotations.Our code is available as a part of an analysis tool with a user interface ready to be used by ecologists.

  • 18.
    Irl, Severin D. H.
    et al.
    Univ Bayreuth, BayCEER, Dept Disturbance Ecol, D-95447 Bayreuth, Germany.
    Steinbauer, Manuel J.
    Univ Bayreuth, BayCEER, Dept Biogeog, D-95447 Bayreuth, Germany.
    Messinger, Jana
    Univ Bayreuth, BayCEER, Dept Biogeog, D-95447 Bayreuth, Germany.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Univ Bayreuth, BayCEER, Dept Biogeog, D-95447 Bayreuth, Germany.
    Palomares-Martinez, Angel
    Parque Nacl Caldera Taburiente, El Paso 38750, Spain.
    Beierkuhnlein, Carl
    Univ Bayreuth, BayCEER, Dept Biogeog, D-95447 Bayreuth, Germany.
    Jentsch, Anke
    Univ Bayreuth, BayCEER, Dept Disturbance Ecol, D-95447 Bayreuth, Germany.
    Burned and devoured-Introduced herbivores, fire, and the endemic flora of the high-elevation ecosystem on La Palma, Canary Islands2014In: Arctic, Antarctic and Alpine research, ISSN 1523-0430, E-ISSN 1938-4246, Vol. 46, no 4, p. 859-869Article in journal (Refereed)
    Abstract [en]

    Novel disturbance regimes (e.g., introduced herbivores and fire) are among the major drivers of degradation in island ecosystems. High-elevation ecosystems (HEEs) on islands might be especially vulnerable to these disturbances due to high endemism. Here, data from an 11-year exclosure experiment in the HEE of La Palma (Canary Islands) are presented where mammalian herbivores have been introduced. We investigate the combined effect of herbivory and fire on total species richness, seedling richness, and seedling establishment on the whole system and a subset of highly endangered species (target species). Total species richness, seedling species richness, and seedling establishment decreased with herbivory. Five out of eight target species were exclusively found inside the exclosures indicating the negative impact of introduced herbivores on endemic high elevation flora. Target species were generally affected more negatively by introduced herbivores and were subject to significantly higher browsing pressure, probably owing to their lack of defense strategies. A natural wildfire that occurred six years before data sampling substantially increased total species richness and seedling richness in both herbivory exclosure and reference conditions. We conclude that species composition of the HEE has been severely altered by the introduction of non-native herbivores, even though fire seems to have a positive effect on this system.

  • 19.
    Jonsson, Hanna
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Olofsson, Johan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Klaminder, Jonatan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Cascading effects of earthworm invasion increase graminoid density and rodent grazing intensities2023In: Ecology, ISSN 0012-9658, E-ISSN 1939-9170Article in journal (Refereed)
    Abstract [en]

    Human-mediated dispersal of non-native earthworms can cause substantial changes to the functioning and composition of ecosystems previously earthworm-free. Some of these earthworm species have the potential to “geoengineer” soils and increase plant nitrogen (N) uptake. Yet the possible consequences of increased plant N concentrations on rodent grazing remains poorly understood. In this study, we present findings from a common garden experiment with two tundra communities, meadow (forb dominated) and heath (shrub dominated), half of them subjected to 4 years of earthworm presence (Lumbricus spp. and Aporrectodea spp.). Within four summers, our earthworm treatment changed plant community composition by increasing graminoid density by, on average, 94% in the heath vegetation and by 49% in the meadow. Rodent winter grazing was more intense on plants growing in soils with earthworms, an effect that coincided with higher N concentrations in plants, indicating a higher palatability. Even though earthworms reduced soil moisture, plant community productivity, as indicated by vegetation greenness (normalized difference vegetation index), was not negatively impacted. We conclude that earthworm-induced changes in plant composition and trophic interactions may fundamentally alter the functioning of tundra ecosystems.

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  • 20. Jurasinski, Gerald
    et al.
    Ahmad, Sate
    Anadon-Rosell, Alba
    Berendt, Jacqueline
    Beyer, Florian
    Bill, Ralf
    Blume-Werry, Gesche
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, 17487 Greifswald, Germany.
    Couwenberg, John
    Guenther, Anke
    Joosten, Hans
    Koebsch, Franziska
    Köhn, Daniel
    Koldrack, Nils
    Kreyling, Jürgen
    Leinweber, Peter
    Lennartz, Bernd
    Liu, Haojie
    Michaelis, Dierk
    Mrotzek, Almut
    Negassa, Wakene
    Schenk, Sandra
    Schmacka, Franziska
    Schwieger, Sarah
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, 17487 Greifswald, Germany.
    Smiljanic, Marko
    Tanneberger, Franziska
    Teuber, Laurenz M.
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, 17487 Greifswald, Germany.
    Urich, Tim
    Wang, Haitao
    Weil, Micha
    Wilmking, Martin
    Zak, Dominik
    Wrage-Monnig, Nicole
    From Understanding to Sustainable Use of Peatlands: The WETSCAPES Approach2020In: SOIL SYSTEMS, ISSN 2571-8789, Vol. 4, no 1, article id 14Article in journal (Refereed)
    Abstract [en]

    Of all terrestrial ecosystems, peatlands store carbon most effectively in long-term scales of millennia. However, many peatlands have been drained for peat extraction or agricultural use. This converts peatlands from sinks to sources of carbon, causing approx. 5% of the anthropogenic greenhouse effect and additional negative effects on other ecosystem services. Rewetting peatlands can mitigate climate change and may be combined with management in the form of paludiculture. Rewetted peatlands, however, do not equal their pristine ancestors and their ecological functioning is not understood. This holds true especially for groundwater-fed fens. Their functioning results from manifold interactions and can only be understood following an integrative approach of many relevant fields of science, which we merge in the interdisciplinary project WETSCAPES. Here, we address interactions among water transport and chemistry, primary production, peat formation, matter transformation and transport, microbial community, and greenhouse gas exchange using state of the art methods. We record data on six study sites spread across three common fen types (Alder forest, percolation fen, and coastal fen), each in drained and rewetted states. First results revealed that indicators reflecting more long-term effects like vegetation and soil chemistry showed a stronger differentiation between drained and rewetted states than variables with a more immediate reaction to environmental change, like greenhouse gas (GHG) emissions. Variations in microbial community composition explained differences in soil chemical data as well as vegetation composition and GHG exchange. We show the importance of developing an integrative understanding of managed fen peatlands and their ecosystem functioning. 

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  • 21.
    Keuper, Frida
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. BioEcoAgro Joint Research Unit, INRAE, Barenton-Bugny, France.
    Wild, Birgit
    Kummu, Matti
    Beer, Christian
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Fontaine, Sébastien
    Gavazov, Konstantin
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Swiss Federal Institute for Forest, Snow and Landscape Research (WSL), Lausanne, Switzerland.
    Gentsch, Norman
    Guggenberger, Georg
    Hugelius, Gustaf
    Jalava, Mika
    Koven, Charles
    Krab, Eveline J.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Soil and Environment, Swedish University of Agricultural Sciences, Uppsala, Sweden.
    Kuhry, Peter
    Monteux, Sylvain
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Richter, Andreas
    Shahzad, Tanvir
    Weedon, James T.
    Dorrepaal, Ellen
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Carbon loss from northern circumpolar permafrost soils amplified by rhizosphere priming2020In: Nature Geoscience, ISSN 1752-0894, E-ISSN 1752-0908, Vol. 13, no 8, p. 560-565Article in journal (Refereed)
    Abstract [en]

    As global temperatures continue to rise, a key uncertainty of climate projections is the microbial decomposition of vast organic carbon stocks in thawing permafrost soils. Decomposition rates can accelerate up to fourfold in the presence of plant roots, and this mechanism—termed the rhizosphere priming effect—may be especially relevant to thawing permafrost soils as rising temperatures also stimulate plant productivity in the Arctic. However, priming is currently not explicitly included in any model projections of future carbon losses from the permafrost area. Here, we combine high-resolution spatial and depth-resolved datasets of key plant and permafrost properties with empirical relationships of priming effects from living plants on microbial respiration. We show that rhizosphere priming amplifies overall soil respiration in permafrost-affected ecosystems by ~12%, which translates to a priming-induced absolute loss of ~40 Pg soil carbon from the northern permafrost area by 2100. Our findings highlight the need to include fine-scale ecological interactions in order to accurately predict large-scale greenhouse gas emissions, and suggest even tighter restrictions on the estimated 200 Pg anthropogenic carbon emission budget to keep global warming below 1.5 °C.

  • 22.
    Krab, Eveline J.
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Soil and Environment, Swedish University of Agricultural Sciences, Uppsala, Sweden.
    Roennefarth, Jonas
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Becher, Marina
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Keuper, Frida
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. INRA, AgroImpact UR1158, Barenton Bugny, France.
    Klaminder, Jonatan
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Kreyling, Juergen
    Makoto, Kobayashi
    Milbau, Ann
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Department of Biodiversity and Natural Environment, Research Institute for Nature and Forest - INBO, Brussels, Belgium.
    Dorrepaal, Ellen
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Winter warming effects on tundra shrub performance are species-specific and dependent on spring conditions2018In: Journal of Ecology, ISSN 0022-0477, E-ISSN 1365-2745, Vol. 106, no 2, p. 599-612Article in journal (Refereed)
    Abstract [en]

    Climate change-driven increases in winter temperatures positively affect conditions for shrub growth in arctic tundra by decreasing plant frost damage and stimulation of nutrient availability. However, the extent to which shrubs may benefit from these conditions may be strongly dependent on the following spring climate. Species-specific differences in phenology and spring frost sensitivity likely affect shrub growth responses to warming. Additionally, effects of changes in winter and spring climate may differ over small spatial scales, as shrub growth may be dependent on natural variation in snow cover, shrub density and cryoturbation. We investigated the effects of winter warming and altered spring climate on growing-season performance of three common and widespread shrub species in cryoturbated non-sorted circle arctic tundra. By insulating sparsely vegetated non-sorted circles and parts of the surrounding heath with additional snow or gardening fleeces, we created two climate change scenarios: snow addition increased soil temperatures in autumn and winter and delayed snowmelt timing without increasing spring temperatures, whereas fleeces increased soil temperature similarly in autumn and winter, but created warmer spring conditions without altering snowmelt timing. Winter warming affected shrub performance, but the direction and magnitude were species-specific and dependent on spring conditions. Spring warming advanced, and later snowmelt delayed canopy green-up. The fleece treatment did not affect shoot growth and biomass in any shrub species despite decreasing leaf frost damage in Empetrum nigrum. Snow addition decreased frost damage and stimulated growth of Vaccinium vitis-idaea by c. 50%, while decreasing Betula nana growth (p < .1). All of these effects were consistent the mostly barren circles and surrounding heath. Synthesis. In cryoturbated arctic tundra, growth of Vaccinium vitis-idaea may substantially increase when a thicker snow cover delays snowmelt, whereas in longer term, warmer winters and springs may favour E. nigrum instead. This may affect shrub community composition and cover, with potentially far-reaching effects on arctic ecosystem functioning via its effects on cryoturbation, carbon cycling and trophic cascading. Our results highlight the importance of disentangling effects of winter and spring climate change timing and nature, as spring conditions are a crucial factor in determining the impact of winter warming on plant performance.

  • 23.
    Makoto, Kobayashi
    et al.
    Teshio Experimental Forest, Field Science Center for Northern Biosphere, Hokkaido University, Horonobe, Japan.
    Kitagawa, Ryo
    Kansai Research Center, Forestry and Forest Products Research Institute, Kyoto, Japan.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    How do leaf functional traits and age influence the maximum rooting depth of trees?2023In: European Journal of Forest Research, ISSN 1612-4669, E-ISSN 1612-4677, Vol. 142, no 5, p. 1197-1206Article in journal (Refereed)
    Abstract [en]

    Maximum rooting depth is a key functional trait to increase the fitness of trees and also influences terrestrial ecosystem processes. Despite its importance, the drivers of the interspecific variation of maximum rooting depth or its relation to other plant traits and plant age are not well understood. In this study, we aimed to clarify the drivers of the interspecific variation of maximum rooting depth with special reference to its relation to plant leaf traits and age. We analyzed how maximum rooting depth of single individuals of 227 tree species planted in the same common garden in the temperate region of central Japan is correlated to commonly measured leaf functional traits (specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen (N) concentration) extracted from the TRY database. We did this by employing the phylogenetic comparable method and included the age of all target trees. When excluding the effect of phylogenetic signals from the relationships between rooting depth and leaf traits, SLA was negatively correlated with maximum rooting depth in deciduous, but not evergreen species. Further, rooting depth and leaf N concentration were negatively correlated in evergreen trees, a pattern driven by young trees. These results implicate that the relationship between maximum rooting depth and leaf traits differed depending on the leaf habits and age of the tree species.

  • 24.
    Malyshev, Andrey V.
    et al.
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Spiller, Ophelia
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Smiljanić, Marko
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Weigel, Robert
    Plant Ecology and Ecosystems Research, University of Goettingen, Göttingen, Germany; Ecological-Botanical Garden, University of Bayreuth, Bayreuth, Germany.
    Kolb, Alexander
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Nze, Byron Ye
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Märker, Frederik
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Sommer, Freymuth Carl-Fried Johannes
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Kinley, Kinley
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany; Ecological-Botanical Garden, University of Bayreuth, Bayreuth, Germany.
    Ziegler, Jan
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany; Swiss Federal Research Institute WSL, Birmensdorf, Switzerland.
    Pasang, Pasang
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Mahara, Robert
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Joshi, Silviya
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Heinsohn, Vincent
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Kreyling, Juergen
    Institute of Botany and Landscape Ecology, University of Greifswald, Greifswald, Germany.
    Warming nondormant tree roots advances aboveground spring phenology in temperate trees2023In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 240, no 6, p. 2276-2287Article in journal (Refereed)
    Abstract [en]

    Climate warming advances the onset of tree growth in spring, but above- and belowground phenology are not always synchronized. These differences in growth responses may result from differences in root and bud dormancy dynamics, but root dormancy is largely unexplored. We measured dormancy in roots and leaf buds of Fagus sylvatica and Populus nigra by quantifying the warming sum required to initiate above- and belowground growth in October, January and February. We furthermore carried out seven experiments, manipulating only the soil and not air temperature before or during tree leaf-out to evaluate the potential of warmer roots to influence budburst timing using seedlings and adult trees of F. sylvatica and seedlings of Betula pendula. Root dormancy was virtually absent in comparison with the much deeper winter bud dormancy. Roots were able to start growing immediately as soils were warmed during the winter. Interestingly, higher soil temperature advanced budburst across all experiments, with soil temperature possibly accounting for c. 44% of the effect of air temperature in advancing aboveground spring phenology per growing degree hour. Therefore, differences in root and bud dormancy dynamics, together with their interaction, likely explain the nonsynchronized above- and belowground plant growth responses to climate warming.

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  • 25.
    Monteux, Sylvain
    et al.
    Department of Environmental Science, Stockholm University, Stockholm, Sweden; Bolin Centre for Climate Research, Stockholm University, Stockholm, Sweden; UiT The Arctic University Museum of Norway, Tromsø, Norway.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Gavazov, Konstantin
    Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Birmensdorf, Switzerland.
    Kirchhoff, Leah
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Krab, Eveline J.
    Department of Soil and Environment, Swedish University for Agricultural Sciences SLU, Uppsala, Sweden.
    Lett, Signe
    Department of Biology, University of Copenhagen, Copenhagen, Denmark.
    Pedersen, Emily P.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Swedish Polar Research Secretariat, Abisko Scientific Research Station, Abisko, Sweden.
    Väisänen, Maria
    Ecology and Genetics Research Unit, University of Oulu, Oulu, Finland.
    Controlling biases in targeted plant removal experiments2023In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137Article in journal (Refereed)
    Abstract [en]

    Targeted removal experiments are a powerful tool to assess the effects of plant species or (functional) groups on ecosystem functions. However, removing plant biomass in itself can bias the observed responses. This bias is commonly addressed by waiting until ecosystem recovery, but this is inherently based on unverified proxies or anecdotal evidence. Statistical control methods are efficient, but restricted in scope by underlying assumptions.

    We propose accounting for such biases within the experimental design, using a gradient of biomass removal controls. We demonstrate the relevance of this design by presenting (1) conceptual examples of suspected biases and (2) how to observe and control for these biases.

    Using data from a mycorrhizal association-based removal experiment, we show that ignoring biomass removal biases (including by assuming ecosystem recovery) can lead to incorrect, or even contrary conclusions (e.g. false positive and false negative). Our gradient design can prevent such incorrect interpretations, regardless of whether aboveground biomass has fully recovered.

    Our approach provides more objective and quantitative insights, independently assessed for each variable, than using a proxy to assume ecosystem recovery. Our approach circumvents the strict statistical assumptions of, for example, ANCOVA and thus offers greater flexibility in data analysis.

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  • 26.
    Monteux, Sylvain
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Weedon, James T.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Gavazov, Konstantin
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Federal Institute for Forest, Snow and Landscape Research WSL, Lausanne, Switzerland.
    Jassey, Vincent E. J.
    Johansson, Margareta
    Keuper, Frida
    Olid, Carolina
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Dorrepaal, Ellen
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Long-term in situ permafrost thaw effects on bacterial communities and potential aerobic respiration2018In: The ISME Journal, ISSN 1751-7362, E-ISSN 1751-7370, Vol. 12, no 9, p. 2129-2141Article in journal (Refereed)
    Abstract [en]

    The decomposition of large stocks of soil organic carbon in thawing permafrost might depend on more than climate change-induced temperature increases: indirect effects of thawing via altered bacterial community structure (BCS) or rooting patterns are largely unexplored. We used a 10-year in situ permafrost thaw experiment and aerobic incubations to investigate alterations in BCS and potential respiration at different depths, and the extent to which they are related with each other and with root density. Active layer and permafrost BCS strongly differed, and the BCS in formerly frozen soils (below the natural thawfront) converged under induced deep thaw to strongly resemble the active layer BCS, possibly as a result of colonization by overlying microorganisms. Overall, respiration rates decreased with depth and soils showed lower potential respiration when subjected to deeper thaw, which we attributed to gradual labile carbon pool depletion. Despite deeper rooting under induced deep thaw, root density measurements did not improve soil chemistry-based models of potential respiration. However, BCS explained an additional unique portion of variation in respiration, particularly when accounting for differences in organic matter content. Our results suggest that by measuring bacterial community composition, we can improve both our understanding and the modeling of the permafrost carbon feedback.

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  • 27.
    Peters, Bo
    et al.
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, Soldmannstraße 15, Greifswald, Germany.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, Soldmannstraße 15, Greifswald, Germany.
    Gillert, Alexander
    Fraunhofer Institute for Computer Graphics Research IGD, Rostock, Germany.
    Schwieger, Sarah
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, Soldmannstraße 15, Greifswald, Germany.
    von Lukas, Uwe Freiherr
    Fraunhofer Institute for Computer Graphics Research IGD, Rostock, Germany; Institute for Visual & Analytic Computing, University of Rostock, Rostock, Germany.
    Kreyling, Juergen
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, University of Greifswald, Soldmannstraße 15, Greifswald, Germany.
    As good as human experts in detecting plant roots in minirhizotron images but efficient and reproducible: the convolutional neural network “RootDetector”2023In: Scientific Reports, E-ISSN 2045-2322, Vol. 13, no 1, article id 1399Article in journal (Refereed)
    Abstract [en]

    Plant roots influence many ecological and biogeochemical processes, such as carbon, water and nutrient cycling. Because of difficult accessibility, knowledge on plant root growth dynamics in field conditions, however, is fragmentary at best. Minirhizotrons, i.e. transparent tubes placed in the substrate into which specialized cameras or circular scanners are inserted, facilitate the capture of high-resolution images of root dynamics at the soil-tube interface with little to no disturbance after the initial installation. Their use, especially in field studies with multiple species and heterogeneous substrates, though, is limited by the amount of work that subsequent manual tracing of roots in the images requires. Furthermore, the reproducibility and objectivity of manual root detection is questionable. Here, we use a Convolutional Neural Network (CNN) for the automatic detection of roots in minirhizotron images and compare the performance of our RootDetector with human analysts with different levels of expertise. Our minirhizotron data come from various wetlands on organic soils, i.e. highly heterogeneous substrates consisting of dead plant material, often times mainly roots, in various degrees of decomposition. This may be seen as one of the most challenging soil types for root segmentation in minirhizotron images. RootDetector showed a high capability to correctly segment root pixels in minirhizotron images from field observations (F1 = 0.6044; r2 compared to a human expert = 0.99). Reproducibility among humans, however, depended strongly on expertise level, with novices showing drastic variation among individual analysts and annotating on average more than 13-times higher root length/cm2 per image compared to expert analysts. CNNs such as RootDetector provide a reliable and efficient method for the detection of roots and root length in minirhizotron images even from challenging field conditions. Analyses with RootDetector thus save resources, are reproducible and objective, and are as accurate as manual analyses performed by human experts.

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  • 28.
    Sarneel, Judith M.
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Barel, Janna M.
    Aquatic Ecology & Environmental Biology, Faculty of Science, Radboud Institute for Biological and Environmental Sciences, Radboud University Nijmegen, Nijmegen, Netherlands.
    Duddigan, Sarah
    Soil Research Centre and Department of Geography & Environmental Science, University of Reading, Reading, United Kingdom.
    Keuskamp, Joost A.
    Ecology & Biodiversity Group, Institute of Environmental Biology, Utrecht University, Utrecht, Netherlands; Biont Research, Utrecht, Netherlands.
    Pastor, Ada
    GRECO, Institute of Aquatic Ecology, University of Girona, Girona, Spain.
    Sandén, Taru
    Department for Soil Health and Plant Nutrition, Austrian Agency for Health and Food Safety (AGES), Vienna, Austria.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Reasons to not correct for leaching in TBI; Reply to Lind et al. (2022)2023In: Ecology and Evolution, E-ISSN 2045-7758, Vol. 13, no 6, article id e10133Article in journal (Other academic)
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  • 29.
    Schwieger, Sarah
    et al.
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Germany.
    Blume-Werry, Gesche
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Germany.
    Ciesiolka, Felix
    Anadon-Rosell, Alba
    Root biomass and root traits of Alnus glutinosa show size-dependent and opposite patterns in a drained and a rewetted forest peatland2021In: Annals of Botany, ISSN 0305-7364, E-ISSN 1095-8290, Vol. 127, no 3, p. 337-346Article in journal (Refereed)
    Abstract [en]

    BACKGROUND AND AIMS: Forest peatlands represent 25 % of global peatlands and store large amounts of carbon (C) as peat. Traditionally they have been drained in order to increase forestry yield, which may cause large losses of C from the peat. Rewetting aims to stop these losses and to restore the initial storage function of the peatlands. As roots represent major peat-forming elements in these systems, we sampled roots with diameter <5 mm in a drained and a rewetted forest peatland in north-east Germany to evaluate differences in tree biomass investments below ground, root functional characteristics and root age.

    METHODS: We cored soil next to Alnus glutinosa stems and sorted root biomass into <1, 1-2 and 2-5 mm diameter classes. We measured biomass distribution and specific root area (SRA) in 10-cm depth increments down to 50 cm, and estimated root age from annual growth rings.

    KEY RESULTS: Root biomass in the rewetted site was more than double that in the drained site. This difference was mostly driven by very fine roots <1 mm, which accounted for 51 % of the total root biomass and were mostly (75 %) located in the upper 20 cm. For roots <1 mm, SRA did not differ between the sites. However, SRA of the 1-2 mm and 2-5 mm diameter roots was higher in the drained than in the rewetted site. Root age did not differ between sites.

    CONCLUSIONS: The size-dependent opposite patterns between root biomass and their functional characteristics under contrasting water regimes indicate differences between fine and coarse roots in their response to environmental changes. Root age distribution points to similar root turnover rates between the sites, while higher root biomass in the rewetted site clearly indicates larger tree C stocks below ground under rewetting, supporting the C sink function of the ecosystem.

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  • 30.
    Schwieger, Sarah
    et al.
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Blume-Werry, Gesche
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Peters, Bo
    Smiljanic, Marko
    Kreyling, Juergen
    Patterns and drivers in spring and autumn phenology differ above- and belowground in four ecosystems under the same macroclimatic conditions2019In: Plant and Soil, ISSN 0032-079X, E-ISSN 1573-5036, Vol. 445, no 1-2, p. 217-229Article in journal (Refereed)
    Abstract [en]

    Background and aims: Start and end of the growing season determine important ecosystem processes, but their drivers may differ above-and belowground, between autumn and spring, and between ecosystems.

    Here, we compare above-and belowground spring and autumn phenology, and their abiotic drivers (temperature, water level, and soil moisture) in four temperate ecosystems (beech forest, alder carr, phragmites reed, and sedge reed).

    Methods: Root growth was measured in-situ with minirhizotrons and compared with aboveground phenology assessed with dendrometer data and NDVI.

    Results: Synchrony of above- and belowground phenology depended on ecosystem. Onset of root growth was later than shoot growth in all three peatlands (12–33 days), but similar in the beech forest. The growing season ended earlier belowground in the two forested ecosystems (beech forest: 27 days, understory of the alder carr: 55 days), but did not differ in the phragmites reed. Generally, root production was correlated with soil temperature (especially in spring) and water level in the peatlands, while abiotic factors were less correlated with leaf activity or root production in either spring or autumn in the beech forest.

    Conclusions: Root production on organic soils was ten times higher compared to the zonal broadleaf deciduous forest on mineral soils, highlighting the importance of peatlands. Belowground phenology cannot be projected from aboveground phenology and measuring root phenology is crucial to understand temporal dynamics of production and carbon fluxes.

  • 31.
    Schwieger, Sarah
    et al.
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Germany.
    Kreyling, Juergen
    Couwenberg, John
    Smiljanic, Marko
    Weigel, Robert
    Wilmking, Martin
    Blume-Werry, Gesche
    Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Germany.
    Wetter is Better: Rewetting of Minerotrophic Peatlands Increases Plant Production and Moves Them Towards Carbon Sinks in a Dry Year2021In: Ecosystems (New York. Print), ISSN 1432-9840, E-ISSN 1435-0629, Vol. 24, no 5, p. 1093-1109Article in journal (Refereed)
    Abstract [en]

    Peatlands are effective carbon sinks as more biomass is produced than decomposed under the prevalent anoxic conditions. Draining peatlands coupled with warming releases stored carbon, and subsequent rewetting may or may not restore the original carbon sink. Yet, patterns of plant production and decomposition in rewetted peatlands and how they compare to drained conditions remain largely unexplored. Here, we measured annual above- and belowground biomass production and decomposition in three different drained and rewetted peatland types: alder forest, percolation fen and coastal fen during an exceptionally dry year. We also used standard plant material to compare decomposition between the sites, regardless of the decomposability of the local plant material. Rewetted sites showed higher root and shoot production in the percolation fen and higher root production in the coastal fen, but similar root and leaf production in the alder forest. Decomposition rates were generally similar in drained and rewetted sites, only in the percolation fen and alder forest did aboveground litter decompose faster in the drained sites. The rewetted percolation fen and the two coastal sites had the highest projected potential for organic matter accumulation. Roots accounted for 23–66% of total biomass production, and belowground biomass, rather than aboveground biomass, was particularly important for organic matter accumulation in the coastal fens. This highlights the significance of roots as main peat-forming element in these graminoid-dominated fen peatlands and their crucial role in carbon cycling, and shows that high biomass production supported the peatlands’ function as carbon sink even during a dry year.

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  • 32. Schwieger, Sarah
    et al.
    Kreyling, Juergen
    Milbau, Ann
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    Autumnal warming does not change root phenology in two contrasting vegetation types of subarctic tundra2018In: Plant and Soil, ISSN 0032-079X, E-ISSN 1573-5036, Vol. 424, no 1-2, p. 145-156Article in journal (Refereed)
    Abstract [en]

    Root phenology is important in controlling carbon and nutrient fluxes in terrestrial ecosystems, yet, remains largely unexplored, especially in the Arctic. We compared below- and aboveground phenology and ending of the growing season in two contrasting vegetation types of subarctic tundra: heath and meadow, and their response to experimental warming in autumn. Root phenology was measured in-situ with minirhizotrons and compared with aboveground phenology assessed with repeat digital photography. The end of the growing season, both below- and aboveground, was similar in meadow and heath and the belowground growing season ended later than aboveground in the two vegetation types. Root growth was higher and less equally distributed over time in meadow compared to heath. The warming treatment increased air and soil temperature by 0.5 A degrees C and slightly increased aboveground greenness, but did not affect root growth or prolong the below- and aboveground growing season in either of the vegetation types. These results imply that vegetation types differ in root dynamics and suggest that other factors than temperature control autumnal root growth in these ecosystems. Further investigations of root phenology will help to identify those drivers, in which including responses of functionally contrasting vegetation types will help to estimate how climate change affects belowground processes and their roles in ecosystem function.

  • 33.
    Schwieger, Sarah
    et al.
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology, Institute of Botany and Landscape Ecology, Greifswald University, Greifswald, Germany.
    Kreyling, Juergen
    Experimental Plant Ecology Institute of Botany and Landscape Ecology, Greifswald University Greifswald Germany.
    Peters, Bo
    Experimental Plant Ecology Institute of Botany and Landscape Ecology, Greifswald University Greifswald Germany.
    Gillert, Alexander
    Fraunhofer Institute for Computer Graphics Research IGD Rostock Germany.
    Freiherr von Lukas, Uwe
    Fraunhofer Institute for Computer Graphics Research IGD Rostock Germany.
    Jurasinski, Gerald
    Faculty of Agriculture and Environmental Sciences University of Rostock Rostock Germany.
    Köhn, Daniel
    Faculty of Agriculture and Environmental Sciences University of Rostock Rostock Germany.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences. Experimental Plant Ecology Institute of Botany and Landscape Ecology, Greifswald University.
    Rewetting prolongs root growing season in minerotrophic peatlands and mitigates negative drought effects2022In: Journal of Applied Ecology, ISSN 0021-8901, E-ISSN 1365-2664, Vol. 59, no 8, p. 2106-2116Article in journal (Refereed)
    Abstract [en]

    Root phenology influences the timing of plant resource acquisition and carbon fluxes into the soil. This is particularly important in fen peatlands, in which peat is primarily formed by roots and rhizomes of vascular plants. However, most fens in Central Europe are drained for agriculture, leading to large carbon losses, and further threatened by increasing frequency and intensity of droughts. Rewetting fens aims to restore the original carbon sink, but how root phenology is affected by drainage and rewetting is largely unknown.

    We monitored root phenology with minirhizotrons in drained and rewetted fens (alder forest, percolation fen and coastal fen) as well as its soil temperature and water table depth during the 2018 drought. For each fen type, we studied a drained site and a site that was rewetted ~25 years ago, while all the sites studied had been drained for almost a century.

    Overall, the growing season was longer with rewetting, allowing roots to grow over a longer period in the year and have a higher root production than under drainage. With increasing depth, the growing season shifted to later in time but remained a similar length, and the relative importance of soil temperature for root length changes increased with soil depth.

    Synthesis and applications: Rewetting extended the growing season of roots, highlighting the importance of phenology in explaining root productivity in peatlands. A longer growing season allows a longer period of carbon sequestration in form of root biomass and promotes the peatlands' carbon sink function, especially through longer growth in deep soil layers. Thus, management practices that focus on rewetting peatland ecosystems are necessary to maintain their function as carbon sinks, particularly under drought conditions, and are a top priority to reduce carbon emissions and address climate change.

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  • 34.
    Spitzer, Clydecia M.
    et al.
    Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, Skogsmarksgränd, Umeå, Sweden.
    Blume-Werry, Gesche
    Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.
    As a permafrost ecosystem warms, plant community traits become more acquisitive2023In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 240, no 5, p. 1712-1713Article in journal (Other academic)
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