Biomechanical traits of salt marsh vegetation are insensitive to future climate scenarios.

Salt marshes provide wave and flow attenuation, making them attractive for coastal protection. It is necessary to predict their coastal protection capacity in the future, when climate change will increase hydrodynamic forcing and environmental parameters such as water temperature and CO2 content. We exposed the European salt marsh species Spartina anglica and Elymus athericus to enhanced water temperature (+ 3°) and CO2 (800 ppm) levels in a mesocosm experiment for 13 weeks in a full factorial design. Afterwards, the effect on biomechanic vegetation traits was assessed. These traits affect the interaction of vegetation with hydrodynamic forcing, forming the basis for wave and flow attenuation. Elymus athericus did not respond to any of the treatments suggesting that it is insensitive to such future climate changes. Spartina anglica showed an increase in diameter and flexural rigidity, while Young's bending modulus and breaking force did not differ between treatments. Despite some differences between the future climate scenario and present conditions, all values lie within the natural trait ranges for the two species. Consequently, this mesocosm study suggests that the capacity of salt marshes to provide coastal protection is likely to remain constantly high and will only be affected by future changes in hydrodynamic forcing.

Subsurface aeration of tidal wetland soils: Root-system structure and aerenchyma connectivity in Spartina (Poaceae).

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.

Plant-Mediated Rhizosphere Oxygenation in the Native Invasive Salt Marsh Grass Elymus athericus.

In the last decades, the spread of Elymus athericus has caused significant changes to the plant community composition and ecosystem services of European marshes. The distribution of E. athericus was typically limited by soil conditions characteristic for high marshes, such as low flooding frequency and high soil aeration. However, recently the spread of E. athericus has begun to also include low-marsh environments. A high-marsh ecotype and a low-marsh ecotype of E. athericus have been described, where the latter possess habitat-specific phenotypic traits facilitating a better adaption for inhabiting low-marsh areas. In this study, planar optodes were applied to investigate plant-mediated sediment oxygenation in E. athericus, which is a characteristic trait for marsh plants inhabiting frequently flooded environments. Under waterlogged conditions, oxygen (O2) was translocated from aboveground sources to the roots, where it leaked out into the surrounding sediment generating oxic root zones below the sediment surface. Oxic root zones were clearly visible in the optode images, and no differences were found in the O2-leaking capacity between ecotypes. Concentration profiles measured perpendicular to the roots revealed that the radius of the oxic root zones ranged from 0.5 to 2.6 mm measured from the root surface to the bulk anoxic sediment. The variation of oxic root zones was monitored over three consecutive light-dark cycles (12 h/12 h). The O2 concentration of the oxic root zones was markedly reduced in darkness, yet the sediment still remained oxic in the immediate vicinity of the roots. Increased stomatal conductance improving the access to atmospheric O2 as well as photosynthetic O2 production are likely factors facilitating the improved rhizosphere oxygenation during light exposure of the aboveground biomass. E. athericus' capacity to oxygenate its rhizosphere is an inheritable trait that may facilitate its spread into low-marsh areas. Furthermore, this trait makes E. athericus a highly competitive species in marshes facing the effects of accelerated sea-level rise, where waterlogged sediment conditions could become increasingly pronounced.

Plant-Sediment Interactions in Salt Marshes - An Optode Imaging Study of O2, pH, and CO 2 Gradients in the Rhizosphere.

In many wetland plants, belowground transport of O2 via aerenchyma tissue and subsequent O2 loss across root surfaces generates small oxic root zones at depth in the rhizosphere with important consequences for carbon and nutrient cycling. This study demonstrates how roots of the intertidal salt-marsh plant Spartina anglica affect not only O2, but also pH and CO2 dynamics, resulting in distinct gradients of O2, pH, and CO2 in the rhizosphere. A novel planar optode system (VisiSens TD®, PreSens GmbH) was used for taking high-resolution 2D-images of the O2, pH, and CO2 distribution around roots during alternating light-dark cycles. Belowground sediment oxygenation was detected in the immediate vicinity of the roots, resulting in oxic root zones with a 1.7 mm radius from the root surface. CO2 accumulated around the roots, reaching a concentration up to threefold higher than the background concentration, and generally affected a larger area within a radius of 12.6 mm from the root surface. This contributed to a lowering of pH by 0.6 units around the roots. The O2, pH, and CO2 distribution was recorded on the same individual roots over diurnal light cycles in order to investigate the interlinkage between sediment oxygenation and CO2 and pH patterns. In the rhizosphere, oxic root zones showed higher oxygen concentrations during illumination of the aboveground biomass. In darkness, intraspecific differences were observed, where some plants maintained oxic root zones in darkness, while others did not. However, the temporal variation in sediment oxygenation was not reflected in the temporal variations of pH and CO2 around the roots, which were unaffected by changing light conditions at all times. This demonstrates that plant-mediated sediment oxygenation fueling microbial decomposition and chemical oxidation has limited impact on the dynamics of pH and CO2 in S. anglica rhizospheres, which may in turn be controlled by other processes such as root respiration and root exudation.

Dynamics of oxygen and carbon dioxide in rhizospheres of Lobelia dortmanna - a planar optode study of belowground gas exchange between plants and sediment.

Root-mediated CO2 uptake, O2 release and their effects on O2 and CO2 dynamics in the rhizosphere of Lobelia dortmanna were investigated. Novel planar optode technology, imaging CO2 and O2 distribution around single roots, provided insights into the spatiotemporal patterns of gas exchange between roots, sediment and microbial community. In light, O2 release and CO2 uptake were pronounced, resulting in a distinct oxygenated zone (radius: c. 3 mm) and a CO2 -depleted zone (radius: c. 2 mm) around roots. Simultaneously, however, microbial CO2 production was stimulated within a larger zone around the roots (radius: c. 10 mm). This gave rise to a distinct pattern with a CO2 minimum at the root surface and a CO2 maximum c. 2 mm away from the root. In darkness, CO2 uptake ceased, and the CO2 -depleted zone disappeared within 2 h. By contrast, the oxygenated root zone remained even after 8 h, but diminished markedly over time. A tight coupling between photosynthetic processes and the spatiotemporal dynamics of O2 and CO2 in the rhizosphere of Lobelia was demonstrated, and we suggest that O2 -induced stimulation of the microbial community in the sediment increases the supply of inorganic carbon for photosynthesis by building up a CO2 reservoir in the rhizosphere.

Survey of sediment oxygenation in rhizospheres of the saltmarsh grass - Spartina anglica.

Although transport of oxygen via the aerenchyma tissue and subsequent oxygen loss across root surfaces is well-documented for salt marsh grasses, only few studies have measured the oxygenation of sediment surrounding roots and rhizomes. In this study, sediment oxygenation was assessed in situ in rhizospheres of the intertidal salt marsh grass, Spartina anglica - an invading species, vigorously spreading in many wetlands around the world. The rhizospheres of two populations of S. anglica with differing plant morphology growing in different sediment types were investigated in situ using a novel multifiber optode system with 100 oxygen probes. No oxygen was detected inside the rhizospheres at any depth in either location, indicating a limited impact of plant-mediated sediment oxygenation on the bulk anoxic sediment. Subsequent planar optode studies imaging the oxygen content around the roots substantiated these findings showing that sediment oxygenation was present in both locations, but it was confined only to the immediate vicinity of the root tips. The size of the oxic zones surrounding the root tips differed between sediment-types: in S. anglica growing in permeable sandy sediment, oxic root zones extended 1.5mm away from the roots surface compared to only 0.4mm in muddy tidal flat deposit, which had a substantially higher oxygen demand. The oxygen concentration inside the oxic root zones remained stable during continuous light and air-exposure of the aboveground biomass. In comparison, sediment oxygenation generated by burrowing infauna (Hediste diversicolor) showed to be markedly more temporally variability, reaching anoxic conditions multiple times during a 5-h period.

CO2 and inorganic nutrient enrichment affect the performance of a calcifying green alga and its noncalcifying epiphyte.

Ocean acidification studies in the past decade have greatly improved our knowledge of how calcifying organisms respond to increased surface ocean CO2 levels. It has become evident that, for many organisms, nutrient availability is an important factor that influences their physiological responses and competitive interactions with other species. Therefore, we tested how simulated ocean acidification and eutrophication (nitrate and phosphate enrichment) interact to affect the physiology and ecology of a calcifying chlorophyte macroalga (Halimeda opuntia (L.) J.V. Lamouroux) and its common noncalcifying epiphyte (Dictyota sp.) in a 4-week fully crossed multifactorial experiment. Inorganic nutrient enrichment (+NP) had a strong influence on all responses measured with the exception of net calcification. Elevated CO2 alone significantly decreased electron transport rates of the photosynthetic apparatus and resulted in phosphorus limitation in both species, but had no effect on oxygen production or respiration. The combination of CO2 and +NP significantly increased electron transport rates in both species. While +NP alone stimulated H. opuntia growth rates, Dictyota growth was significantly stimulated by nutrient enrichment only at elevated CO2, which led to the highest biomass ratios of Dictyota to Halimeda. Our results suggest that inorganic nutrient enrichment alone stimulates several aspects of H. opuntia physiology, but nutrient enrichment at a CO2 concentration predicted for the end of the century benefits Dictyota sp. and hinders its calcifying basibiont H. opuntia.