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In recent years, important processes controlling ecosystem carbon dynamics have been connected to fine-root exudation of soluble carbon compounds. Root exudation patterns may change depending on plant interactions and plant susceptibility to and recovery from drought. Recent investigations suggest that root exudation tends to increase with stress events, but quantification of the amount of carbon released from roots across soil depths with differing water availability and species interactions are missing.We tested if root exudation rates were negatively correlated with soil water content across soil depths during and after drought. We further tested if species in mixture, often considered to be less stressed under drought, exuded less carbon than species in monospecific environments. Exudates were sampled in a mature Fagus sylvatica L. and Picea abies (L.) Karst. forest at the end of a five-year throughfall exclusion period and again one year after the drought ended. We quantified root exudates and their variation with soil depth for both tree species in monospecific and mixed species zones.Carbon exudation significantly increased in fine roots exposed to dry soils (

Forest soils are crucial for many ecosystem services that rely on soil organic matter (SOM) stability. Carbon allocated to roots and released as exudates to the rhizosphere plays a key role in SOM stabilization. Under periodic drought, elevated root exudation and SOM accumulation have been reported. Yet, whether root exudates control SOM formation and stability in mature forests once the drought ends is largely unknown. We examined whether root exudates from P. abies and F. sylvatica trees relate to SOM formation and stability in soil depth profiles one year following five years of experimental drought (Kroof experiment, Germany). We collected root exudates throughout the rooting zone and combined the data with thermogravimetric analysis of SOM in the rhizosphere and non-rooted soil. We found that the rhizosphere of both species was characterized by stable SOM fractions that did not decrease post-drought, suggesting potential protection of SOM due to rhizodeposition and root exudates. In contrast, stable SOM fractions decreased relative to controls in non-rooted topsoil below P. abies, indicating a loss of stabilized SOM from drought-affected and re-wetted soil. Our measurements provide valuable insights into post-drought SOM formation and mechanisms of SOM stabilization in forest ecosystems under climate change.

There is urgency in optimizing agricultural production in light of climate uncertainty, human population growth and resource limitations. The rhizosphere microbiome is a promising genetic resource, yet it is poorly understood. Additionally, research into rhizosphere microbial community assembly (MCA) lacks a consistent ecological framework that incorporates the dynamics of the occurring interactions. We explore the ecological principles that guide community establishment as they pertain to the rhizosphere while reviewing relevant research and highlighting the gaps in knowledge. We propose a conceptual model for studying the rhizosphere, and argue for higher resolution characterizations of the rhizosphere, under a framework of root and microbial traits that will determine its resulting composition. For this, we borrow concepts from ecological theory to chronologically describe MCA as a function of colonization, distribution and succession under changing plant-imposed filters. We argue that there is a need to consider the temporal and spatial scale of rhizosphere processes in a manner that is relevant to its microbial components. Finally, we discuss strategies for managing the rhizosphere microbiome for agriculture and the remaining gaps in knowledge that impede their application. Synthesis and applications. We aim to initiate a comprehensive conversation on ecological processes and plant and microbial traits that affect rhizosphere assembly. Rhizosphere microbiome research is relatively new and highly multidisciplinary. The use of a shared vocabulary and ecological theory will facilitate the development and advances in this discipline.

Temperate forest ecosystems are exposed to a higher frequency, duration and severity of drought. To promote forest longevity in a changing climate, we require a better understanding of the long-term impacts of repetitive drought events on fine-root dynamics in mature forests. Using minirhizotron methods, we investigated the effect of seasonal drought on fine-root dynamics in single-species and mixed-species arrangements of Fagus sylvatica (European beech) and Picea abies (Norway spruce) by means of a 4-yr-long throughfall-exclusion experiment. Fine-root production of both species decreased under drought. However, this reduction was not evident for P. abies when grown intermixed with F. sylvatica. Throughfall-exclusion prolonged the lifespan of P. abies roots but did not change the lifespan of F. sylvatica roots, except in 2016. Fagus sylvatica responded to drought by reducing fine-root production at specific depths and during roof closure. This is the first study to examine long-term trends in mature forest fine-root dynamics under repetitive drought events. Species-specific fine-root responses to drought have implications for the rate and depth of root-derived organic matter supply to soil. From a root dynamics perspective, intermixing tree species is not beneficial to all species but dampens drought impacts on the belowground productivity of P. abies.

With increasing societal demands for food security and environmental sustainability on land, the question arises: to what extent do synergies and trade-offs exist between soil functions and how can they be measured across Europe? To address this challenge, we followed the functional land management approach and assessed five soil functions: primary productivity, water regulation and purification, climate regulation, soil biodiversity and nutrient cycling. Soil, management and climate data were collected from 94 sites covering 13 countries, five climatic zones and two land-use types (arable and grassland). This dataset was analysed using the Soil Navigator, a multicriteria decision support system developed to assess the supply of the five soil functions simultaneously. Most sites scored high for two to three soil functions, demonstrating that managing for multifunctionality in soil is possible but that local constraints and trade-offs do exist. Nutrient cycling, biodiversity and climate regulation were less frequently delivered at high capacity than the other two soil functions. Using correlation and co-occurrence analyses, we also found that synergies and trade-offs between soil functions vary among climatic zones and land-use types. This study provides a new framework for monitoring soil quality at the European scale where both the supply of soil functions and their interactions are considered. Highlights: Managing and monitoring soil multifunctionality across Europe is possible. Synergies and trade-offs between soil functions exist, making it difficult to maximize the supply of all five soil functions simultaneously. Synergies and trade-offs between soil functions vary by climatic zone and land-use type. Climate regulation, biodiversity and nutrient cycling are less frequently delivered at high capacity.

Soil biota contribute to the delivery of multiple soil functions. However, soil biological methods are highly underrepresented in the assessment of soil functionality in agricultural production systems. Here we present a flexible tool to support decision-making during the selection process of soil biological methods for monitoring soil functions. This tool is based on a structured and conceptual framework that connects soil biota to soil functions through their contribution to different soil processes. The methods assessed by the tool were selected as a result of a thorough literature review. Soil biology experts supported the development of the tool (i) by providing feedback on the reviewed methods through a survey and (ii) by determining the relevance of different soil biota to the soil processes related to soil multifunctionality during a workshop. The tool is freely accessible online at the Biological Soil Information System (BIOSIS) platform, where researchers or users with an understanding of research practices can interact with the tool to define the context of their assessment and preferences for technical criteria of the methods. By incorporating user input, this flexible tool can help inform a wide variety of research and assessment programs interested in applying soil biological methods to monitor soil multifunctionality at different scales.

Carbon (C) exuded via roots is proposed to increase under drought and facilitate important ecosystem functions. However, it is unknown how exudate quantities relate to the total C budget of a drought-stressed tree, that is, how much of net-C assimilation is allocated to exudation at the tree level. - We calculated the proportion of daily C assimilation allocated to root exudation during early summer by collecting root exudates from mature Fagus sylvatica and Picea abies exposed to experimental drought, and combining above- and belowground C fluxes with leaf, stem and fine-root surface area. - Exudation from individual roots increased exponentially with decreasing soil moisture, with the highest increase at the wilting point. Despite c. 50% reduced C assimilation under drought, exudation from fine-root systems was maintained and trees exuded 1.0% (F. sylvatica) to 2.5% (P. abies) of net C into the rhizosphere, increasing the proportion of C allocation to exudates two- to three-fold. Water-limited P. abies released two-thirds of its exudate C into the surface soil, whereas in droughted F. sylvatica it was only one-third. - Across the entire root system, droughted trees maintained exudation similar to controls, suggesting drought-imposed belowground C investment, which could be beneficial for ecosystem resilience.

Drought is a severe natural risk that increases drying-rewetting frequencies of soils. Yet, it remains largely unknown how forest ecosystems respond to dry-wet cycles, hampering our ability to evaluate the overall sink and source functionality for this large carbon pool. Recent investigations suggest that the release of soluble carbon via root exudation increases under drought, influencing soil carbon stabilization and mineralization. However, an integration of root exudation into the carbon allocation dynamics of drought stressed trees is missing. We hypothesized that roots in dry soil layers have a higher exudation rate than roots in more moist layers across different soil depths. Further, we tested if higher exudation rates under drought are attenuated by reduced root abundance in dry soils and if the fraction of root exudation from total carbon allocation increases with decreasing photosynthesis rates under drought. At the KROOF experimental site in southern Germany, where mature beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) trees were exposed to artificial drought stress for five consecutive growing seasons, we show that at the root level root exudation rate increases in drier soils. Especially roots in the upper soil profile and roots of spruce trees increased root exudation under drought. When scaled to whole tree level, we did not find differences in total exudation between drought stressed and control trees, indicating sustained root exudation at the tree level under drought. As photosynthesis rates and therefore total carbon assimilation was substantially reduced under drought (by 50 % in beech and almost 70 % in spruce), the fraction of root exudation from total assimilation slightly increased for drought stressed trees. Our results demonstrate that stimulation of root exudation rates with drought exists in natural temperate forest ecosystems but might be mitigated by reduced fine root abundance under drought. Nevertheless, increased exudation per root surface area will have localized impacts on rhizosphere microbial composition and activity especially in the topsoil exposed to more extreme dry-wet cycles. Finally, also the exudate composition can help to determine how priming of soil organic matter relates to belowground carbon allocation dynamics and to disclose processes of complementary species interaction and should be emphasised in future studies.

In Latin America, the cultivation of Arabica coffee (Coffea arabica) plays a critical role in rural livelihoods, biodiversity conservation, and sustainable development. Over the last 20 years, coffee farms and landscapes across the region have undergone rapid and profound biophysical changes in response to low coffee prices, changing climatic conditions, severe plant pathogen outbreaks, and other drivers. Although these biophysical transformations are pervasive and affect millions of rural livelihoods, there is limited information on the types, location, and extent of landscape changes and their socioeconomic and ecological consequences. Here we review the state of knowledge on the ongoing biophysical changes in coffee-growing regions, explore the potential socioeconomic and ecological impacts of these changes, and highlight key research gaps. We identify seven major land-use trends which are affecting the sustainability of coffee-growing regions across Latin America in different ways. These trends include (1) the widespread shift to disease-resistant cultivars, (2) the conventional intensification of coffee management with greater planting densities, greater use of agrochemicals and less shade, (3) the conversion of coffee to other agricultural land uses, (4) the introduction of Robusta coffee (Coffea canephora) into areas not previously cultivated with coffee, (5) the expansion of coffee into forested areas, (6) the urbanization of coffee landscapes, and (7) the increase in the area of coffee produced under voluntary sustainability standards. Our review highlights the incomplete and scattered information on the drivers, patterns, and outcomes of biophysical changes in coffee landscapes, and lays out a detailed research agenda to address these research gaps and elucidate the effects of different landscape trajectories on rural livelihoods, biodiversity conservation, and other aspects of sustainable development. A better understanding of the drivers, patterns, and consequences of changes in coffee landscapes is vital for informing the design of policies, programs, and incentives for sustainable coffee production. Supplementary Information The online version contains supplementary material available at 10.1007/s13593-021-00712-0.

Phenolic compounds perform various functions in soil ranging from microbial substrate to toxin and form the basis of several plant-mediated processes. The aim of this study was to investigate how phenolics commonly exuded by tree roots influence soil organic matter (SOM) decomposition and interact with other labile forms of carbon (C) abundant in root exudates. Therefore, we performed a 38-day incubation experiment and assessed whether phenolic compounds (benzoic acid, caffeic acid and catechin) facilitated or inhibited SOM decomposition in a glucose-amended forest soil. Changes in decomposition, substrate use, fungal and bacterial community composition, and microbial abundance and activity were measured over time using 13C-stable-isotope tracing, DNA-based molecular methods and enzyme assays. Our findings showed that phenolics inhibited microbial activity and abundance to varying degrees. Yet, benzoic acid was the only compound producing a substantial priming effect leading to a 21% increase in SOM decomposition, which was amplified in glucose-amended soils. This stimulation in microbial activity was associated with an increase in β-1,4-glucosidase activity and the bacterial genera Paraburkholderia and Caballeronia of the Burkholderiaceae family. Phenolics drove microbial community shifts in glucose-amended soils with negligible interactive effects. In conclusion, phenolic priming of SOM decomposition is associated with microbial community shifts and amplified in the presence of glucose. This evidence emphasizes the need for considering phenolics and interactions among root exudates as priming mechanisms in the rhizosphere and other soil environments where aromatics and phenolic compounds are abundant.

Absorptive fine roots are an important driver of soil biogeochemical cycles. Yet, the spatio-temporal dynamics of those roots in the presence of neighboring species remain poorly understood. The aim of this study was to analyze shifts in absorptive fine-root traits in monoculture or mixtures of Fagus sylvatica [L.] and Picea abies [L.] Karst. We hypothesized that root competition would be higher under single-species than mixed-species interactions, leading to changes in (i) root survivorship, diameter and respiration and (ii) spatio-temporal patterns of root growth and death. Using minirhizotron methods, we monitored the timing and location of absorptive fine-root growth and death at an experimental forest in southern Germany from 2011 to 2013. We also measured root respiration in the spring and fall seasons of 2012 and 2013. Our findings show that the absorptive fine roots of F. sylvatica had a 50% higher risk of root mortality and higher respiration rates in the single-species compared to mixed-species zones. These results support our hypothesis that root competition is less intense for F. sylvatica in mixture versus monoculture. We were unable to find confirmation for the same hypothesis for P. abies. To analyze spatio-temporal patterns of absorptive fine-root production and mortality, we used a mixed-effects model considering root depth (space) and seasons (time) simultaneously. This analysis showed that F. sylvatica shifts root production towards shallower soil layers in mixed-species stands, besides significant seasonal fluctuations in root production depths for both species. Ultimately, the impact of neighbor species identity on root traits observed in this study has important implications for where, when and how fast root-facilitated carbon cycling takes place in single-species versus mixed-species forests. In addition, our study highlights the need for inclusion of absorptive fine-root spatio-temporal dynamics when examining belowground plant interactions and biogeochemical cycles.

Root–soil interactions fundamentally affect the terrestrial carbon (C) cycle and thereby ecosystem feedbacks to climate change. This study addressed the question of whether the secondary metabolism of different temperate forest tree species can affect soil microbial respiration. We hypothesized that phenolics can both increase and decrease respiration depending on their function as food source, mobilizer of other soil resources, signaling compound, or toxin. We analyzed the phenolic compounds from root exudates and root tissue extracts of six tree species grown in a glasshouse using high-performance liquid chromatography. We then tested the effect of individual phenolic compounds, representing the major identified phenylpropanoid compound classes, on microbial respiration through a 5-d soil incubation. Phenolic root profiles were highly species-specific. Of the eight classes identified, flavonoids were the most abundant, with flavanols being the predominating sub-class. Phenolic effects on microbial respiration ranged from a 26% decrease to a 46% increase, with reduced respiration occurring in the presence of compounds possessing a catechol ring. Tree species variation in root phenolic composition influences the magnitude and direction of root effects on microbial respiration. Our data support the hypothesis that functional group rather than biosynthetic class determines the root phenolic effect on soil C cycling.

Aims: The objectives of this study were to evaluate (1) the fertilizer potential of bone char, (2) the effects of wood biochar on plant-available phosphorus (P), and (3) the role of root-mycorrhizae-biochar interactions in plant P acquisition from a P-fixing soil. Methods: Incubation and pot experiments were conducted with a P-fixing soil and maize with or without root hairs and arbuscular mycorrhizae (AM) inoculation. Olsen-, resin-P and plant P accumulation were used to estimate P availability from bone char, co-pyrolyzed bone char-wood biochar, and separate bone char and wood biochar additions produced at 60, 350 and 750 °C, and Triple Superphosphate (TSP). Results: Maize inoculated with AM showed similar P accumulation when fertilized with either 750 °C bone char or TSP. Pyrolyzing bone did not increase extractable P in soil in comparison to unpyrolyzed bone, apart from a 67 % increase in resin-extractable P after additions of bone char pyrolyzed at 350 °C. Despite greater Olsen-P extractability, co-pyrolysis of bone with wood reduced maize P uptake. Wood biochars reduced resin-P from bone char by 14–26 %, whereas oven-dried wood increased resin-P by 23 %. Conclusions: Bone char is an effective P fertilizer, especially if root-AM interactions are simultaneously considered. Biochar influences plant access to soil P and requires careful management to improve P availability.

Pyrolysis of slaughterhouse waste could promote more sustainable phosphorus (P) usage through the development of alternative P fertilizers. This study investigated how pyrolysis temperature (220, 350, 550 and 750 °C), rendering before pyrolysis, and wood or corn biomass additions affect P chemistry in bone char, plant availability, and its potential as P fertilizer.Linear combination fitting of synchrotron-based X-ray absorption near edge structure spectra demonstrated that higher pyrolysis temperatures decreased the fit with organic P references, but increased the fit with a hydroxyapatite (HA) reference, used as an indicator of high calcium phosphate (CaP) crystallinity. The fit to the HA reference increased from 0% to 69% in bone with meat residue and from 20% to 95% in rendered bone. Biomass additions to the bone with meat residue reduced the fit to the HA reference by 83% for wood and 95% for corn, and additions to rendered bone by 37% for wood. No detectable aromatic P forms were generated by pyrolysis. High CaP crystallinity was correlated with low water-extractable P, but high formic acid-extractable P indicative of high plant availability. Bone char supplied available P which was only 24% lower than Triple Superphosphate fertilizer and two- to five-fold higher than rock phosphate.Pyrolysis temperature and biomass additions can be used to design P fertilizer characteristics of bone char through changing CaP crystallinity that optimize P availability to plants.