While the metabolic composition of root exudates is potentially decisive for understanding plant–microbe interactions and the role of ecosystems as carbon sinks, technical constraints continue to hinder a comprehensive understanding of the metabolic composition of root exudates across the plant kingdom. To evaluate the current state-of-the-art in untargeted root exudate chemical analysis, we conducted a literature review on untargeted metabolomics of root exudates. Using two key ecological topics, plant developmental stage and drought effects, we identify current knowledge on the functional role of the metabolite composition of root exudates. Our review of 127 experiments from 59 studies highlights reliance on analytical methods such as LC-MS and GC-MS for the chemical identification of root exudates, which revealed an average of 874 metabolites per study. Shikimates, phenylpropanoids, and carbohydrates dominated the metabolite profiles, yet limitations such as considerable methodological variation and incomplete annotation hinder comparability and broader ecological interpretations. We argue that addressing these gaps demands a cohesive framework encompassing representative sampling methods, integration of targeted and untargeted analyses, and advanced bioinformatic tools for comprehensive annotation. Furthermore, contextualising metabolite data within ecological and evolutionary frameworks is critical to linking exudate profiles with their functional roles. By adopting this approach, root exudate research can move beyond descriptive analyses to uncover patterns with predictive ecological significance, ultimately advancing our understanding of plant adaptation and ecosystem processes.

ABSTRACT Whole genome duplication (WGD or polyploidization) events shape plant evolution, altering ecological responses and plant traits, particularly those related to cell and tissue size. We studied genetic diversity and phenotypic plasticity in Spartina populations, focusing on hybrid (Spartina × townsendii) and allopolyploid (S. anglica) cytotypes in Wadden Sea salt marshes. Our results reveal low genetic diversity in both cytotypes and a complex response of plant traits to global change factors (drought, elevated CO2 concentration). While WGD increased stomatal length, plasticity varied between cytotypes, with allopolyploids showing higher plasticity, especially under elevated CO2. Biomass allocation patterns differed between cytotypes under global change conditions, suggesting distinct effects on ecosystem functioning, such as belowground carbon sequestration and cycling. The allopolyploid's comparatively fewer, larger‐diameter stems may affect aboveground ecosystem functions differently, including sediment trapping and the slowing of tidal currents. Despite similar genetic backgrounds, allopolyploids did not consistently exhibit higher plasticity, challenging previous assumptions. Our findings highlight the complex interplay between hybridization, WGD, phenotypic plasticity, and ecosystem responses to global change, emphasizing the importance of considering polyploidization in understanding plant adaptation and evolutionary dynamics.

AbstractThe storage of organic carbon in the soils of salt marshes and other coastal blue carbon ecosystems has gained considerable attention by the scientific community for more than a decade now, while the relevance and mechanisms of soil inorganic carbon accumulation remain poorly understood. Using long‐term annual accretion monitoring over 17 years in N = 50 permanent plots distributed across a 1050‐ha salt‐marsh complex of the European Wadden Sea, we identified clear relationships between salt‐marsh vertical growth rates and the soil densities of inorganic and organic carbon. Specifically, we demonstrate a strong positive correlation between vertical accretion and inorganic carbon density while observing a strong negative correlation between vertical accretion and organic carbon density. This decoupling observed between inorganic and organic soil carbon stocks was governed by plant community composition and associated plant traits, which controlled sedimentation processes.

The current climate crisis is associated with rising sea levels, which raises the concerning prospect of losing coastal ecosystems, such as salt marshes. Where inland migration is impossible, salt marshes will only persist if their vertical accretion exceeds the rate of sea-level rise. Positive vertical accretion is mainly driven by sedimentation, whereas negative vertical accretion is driven by erosion and soil compaction, among others. These processes can be influenced by abiotic and biotic factors. The biotic factors, best described by plant functional traits of the salt-marsh vegetation, are, however, not well understood. We assembled a large dataset of 336 plots with vertical accretion time series and plant abundances and coupled it with trait data from salt marsh species of the German Wadden Sea, covering natural unmanaged, anthropogenic unmanaged, and grazed marshes. By using multiple logistic regression analyses, we studied the effects of plant functional traits and distance to the marsh edge on vertical accretion. Mean vertical accretion was in the range of recent sea level rise, except for plots on elevated grazed marshes. There were, however, pronounced local differences in vertical accretion. Positive accretion increased with distance to marsh edge and increasing leaf and stem roughness, described by specific stem length, canopy height, stem mass, leaf mass and leaf area. Except on grazed marshes, leaf traits contributed more strongly to the explanation of positive accretion than stem traits. Negative accretion by e.g., erosion was facilitated by low specific root length and low root and rhizome mass, i.e., lower anchoring capacity. To better assess coastal resilience to sea level rise, our findings suggest (i) to include these effect traits in models and experimental analyses of salt marsh vertical accretion and (ii) to consider effects of vegetation roughness on accretion in salt marsh management schemes.

Whole genome duplications (WGDs) lead to polyploid specimens and are regarded as major drivers for speciation and diversification in plants. One prevalent problem when studying WGDs is that effects of polyploidization in ancient polyploids cannot be disentangled from the consequences of selective evolutionary forces. Cytotypic differences in distribution, phenotypic appearance and in response to surface elevation (determined by HOF-modeling) were identified in a relatively young taxa-group of a hexaploid F1-hybrid (Spartina× townsendii H. Groves & J. Groves, Poaceae) and its dodecaploid descendent (Spartina anglica C.E. Hubbard, Poaceae) using vegetation assessments (1029 plots; 1 × 1 m2) from the European Wadden Sea mainland salt marshes, including elevational and mean high tidal (MHT) data. While the F1-hybrid was mainly present in the eastern part of the Wadden Sea, its dodecaploid descendent occurred in the entire Wadden Sea area. The Spartina cytotypes differed in phenotypes (median of Spartina cover: hexaploid = 25% vs. dodecaploid = 12%) and in elevational niche-optimum (hexaploid = − 49.5 cm MHT vs. dodecaploid = 8.0 cm MHT). High ploidy levels correlated with establishment success in Spartina along geographic gradients but did not seem to increase the capacity to cope with abiotic severity downwards the elevational gradient in salt marshes.

Root-aerenchyma in wetland plants facilitate transport of oxygen from aboveground sources (atmosphere and photosynthesis) to belowground roots and rhizomes, where oxygen can leak out and oxygenate the otherwise anoxic soils. In salt marshes, the soil oxygenation capacity varies among different Spartina-taxa, but little is known about structural pattern and connectivity of root-aerenchyma that facilitates this gas transport. Both environmental conditions and ploidy level play a role for the root-system morphology. Root-system morphology of polyploid Spartina-taxa was studied, quantifying root-tissue volume and root-aerenchyma volume of hexaploid Spartina alterniflora, Spartina maritima, and Spartina × townsendii as well as dodecaploid Spartina anglica from different habitats. Computed tomography (CT)-scan image analysis was applied to quantify the volume of roots and aerenchyma, and to determine the root-system structure (ratio of aerenchyma to root-tissue volumes) and aerenchyma connectivity. On average, Spartina-roots accounted for 12% (v/v) and root-aerenchyma accounted for 1% (v/v) of the soil volume in the pioneer marsh. About 90% (v/v) of all roots were associated with aerenchyma. Root-system structures of S. × townsendii and S. anglica differed and showed clear responses to habitat conditions, such as flooding regime and redox potential. The development of large well-connected aerenchyma fragments were specifically shown in S. anglica and to a minor extend in S. maritima. Aerenchyma in S. alterniflora and S. × townsendii consisted only of smaller fragments. Spartina-dominated tidal marsh soils show high connectivity with the atmosphere via root-aerenchyma. The high ploidy level in S. anglica comes along with high connectivity in root-aerenchyma.

AbstractTidal wetlands are effective carbon sinks, mitigating climate change through the long‐term removal of atmospheric CO2. Studies along surface‐elevation and thus flooding‐frequency gradients in tidal wetlands are often used to understand the effects of accelerated sea‐level rise on carbon sequestration, a process that is primarily determined by the balance of primary production and microbial decomposition. It has often been hypothesized that rates of microbial decomposition would increase with elevation and associated increases in soil oxygen availability; however, previous studies yield a wide range of outcomes and equivocal results. Our mechanistic understanding of the elevation–decomposition relationship is limited because most effort has been devoted to understanding the terminal steps of the decomposition process. A few studies assessed microbial exo‐enzyme activities (EEAs) as initial and rate‐limiting steps that often reveal important insight into microbial energy and nutrient constraints. The present study assessed EEAs and microbial abundance along a coastal ecotone stretching a flooding gradient from tidal flat to high marsh in the European Wadden Sea. We found that stabilization of exo‐enzymes to mineral sediments leads to high specific EEAs at low substrate concentrations in frequently flooded, sediment‐rich zones of the studied ecotone. We argue that the high background activity of a mineral‐associated enzyme pool provides a stable decomposition matrix in highly dynamic, frequently flooded zones. Furthermore, we demonstrate that microbial communities are less nutrient limited in frequently flooded zones, where inputs of nutrient‐rich marine organic matter are higher. This was reflected in both increasing exo‐enzymatic carbon versus nutrient acquisition and decreasing fungal versus bacterial abundance with increasing flooding frequency. Our findings thereby suggest two previously unrecognized mechanisms that may contribute to stimulated microbial activity despite decreasing oxygen availability in response to accelerated sea‐level rise.

The cordgrass Spartina anglica C.E. Hubbard (Poaceae) is an invasive transformer in many salt marsh ecosystems worldwide. Relatively little is known about the capacity of Spartina to accelerate salt marsh succession and to protect salt marshes against sea level rise. We analyzed long-term changes in vegetation and elevation in mainland salt marshes of the European Wadden Sea in Schleswig-Holstein, Germany, to estimate the impact of non-native Spartina on the geomorphological resistance of salt marshes to sea level rise and on changes in species diversity. From 1989 to 2019, the Spartina-zone shifted and expanded upwards to elevations of the high marsh zone and Spartina increased in frequency in several salt marsh vegetation communities. At sites where Spartina dominated the vegetation already three decades ago, elevation and species diversity increased with a higher rate compared to sites lacking Spartina. The median change rates reached for elevation MHT +8.6 versus +1.5 mm per year, for species richness +3 versus $$\pm$$0 species per three decades, and for evenness +0.04 versus −0.08 per three decades, regarding plots with versus without former Spartina dominance, respectively. Invasion of salt marshes by Spartina and its continued, long-term presence were associated with increased elevation and species diversity in the face of sea level rise.

Tidal wetlands have been increasingly recognized as long-term carbon sinks in recent years. Work on carbon sequestration and decomposition processes in tidal wetlands focused so far mainly on effects of global-change factors such as sea-level rise and increasing temperatures. However, little is known about effects of land use, such as livestock grazing, on organic matter decomposition and ultimately carbon sequestration. The present work aims at understanding the mechanisms by which large herbivores can affect organic matter decomposition in tidal wetlands. This was achieved by studying both direct animal-microbe interactions and indirect animal-plant-microbe interactions in grazed and ungrazed areas of two long-term experimental field sites at the German North Sea coast. We assessed bacterial and fungal gene abundance using quantitative PCR, as well as the activity of microbial exo-enzymes by conducting fluorometric assays. We demonstrate that grazing can have a profound impact on the microbial community structure of tidal wetland soils, by consistently increasing the fungi-to-bacteria ratio by 38-42%, and therefore potentially exerts important control over carbon turnover and sequestration. The observed shift in the microbial community was primarily driven by organic matter source, with higher contributions of recalcitrant autochthonous (terrestrial) vs. easily degradable allochthonous (marine) sources in grazed areas favoring relative fungal abundance. We propose a novel and indirect form of animal-plant-microbe interaction: top-down control of aboveground vegetation structure determines the capacity of allochthonous organic matter trapping during flooding and thus the structure of the microbial community. Furthermore, our data provide the first evidence that grazing slows down microbial exo-enzyme activity and thus decomposition through changes in soil redox chemistry. Activities of enzymes involved in C cycling were reduced by 28-40%, while activities of enzymes involved in N cycling were not consistently affected by grazing. It remains unclear if this is a trampling-driven direct grazing effect, as hypothesized in earlier studies, or if the effect on redox chemistry is plant mediated and thus indirect. This study improves our process-level understanding of how grazing can affect the microbial ecology and biogeochemistry of semi-terrestrial ecosystems that may help explain and predict differences in C turnover and sequestration rates between grazed and ungrazed systems.

The rate of biological decomposition greatly determines the carbon sequestration capacity of salt marshes. Microorganisms are involved in the decomposition of biomass and the rate of decomposition is supposed to be related to microbial abundance. Recent studies quantified microbial abundance by means of quantitative polymerase chain reaction (qPCR), a method that also allows determining the microbial community structure by applying specific primers. The main microbial community structure can be determined by using primers specific for 16S rRNA (Bacteria) and 18S rRNA (Fungi) of the microbial DNA. However, the investigation of microbial abundance pattern at small spatial scales, such as locally varying abiotic conditions within a salt-marsh system, requires high accuracy in DNA extraction and qPCR methods. Furthermore, there is evidence that a single extraction may not be sufficient to reliably quantify rRNA gene copies. The aim of this study was to establish a suitable DNA extraction method and stable qPCR conditions for the measurement of microbial abundances in semi-terrestrial environments.