Vegetation zones as indicators of denitrification potential in salt marshes.

Salt marsh vegetation zones shift in response to large-scale environmental changes such as sea-level rise (SLR) and restoration activities, but it is unclear if they are good indicators of soil nitrogen removal. Our goal was to characterize the relationship between denitrification potential and salt marsh vegetation zones in tidally restored and tidally unrestricted coastal marshes, and to use vegetation zones to extrapolate how SLR may influence high marsh denitrification at the landscape scale. We conducted denitrification enzyme activity assays on sediment collected from three vegetation zones expected to shift in distribution due to SLR and tidal flow restoration across 20 salt marshes in Connecticut, USA (n = 60 sampling plots) during the summer of 2017. We found lower denitrification potential in short-form Spartina alterniflora zones (mean, 95% CI: 4, 3-6 mg N h-1  m-2 ) than in S. patens (25, 15-36 mg N h-1  m-2 ) and Phragmites australis (56, 16-96 mg N h-1  m-2 ) zones. Vegetation zone was the single best predictor and explained 52% of the variation in denitrification potential; incorporating restoration status and soil characteristics (soil salinity, moisture, and ammonium) did not improve model fit. Because denitrification potential did not differ between tidally restored and unrestricted marshes, we suggest landscape-scale changes in denitrification after tidal restoration are likely to be associated with shifts in vegetation, rather than differences driven by restoration status. Sea-level-rise-induced hydrologic changes are widely observed to shift high marsh dominated by S. patens to short-form S. alterniflora. To explore the implications of this shift in dominant high marsh vegetation, we paired our measured mean denitrification potential rates with projections of high marsh loss from SLR. We found that, under low and medium SLR scenarios, predicted losses of denitrification potential due to replacement of S. patens by short-form S. alterniflora were substantially larger than losses due to reduced high marsh land area alone. Our results suggest that changes in vegetation zones can serve as landscape-scale predictors of the response of denitrification rates to rapid changes occurring in salt marshes.

Plant-available soil nutrients have a limited influence on cone production patterns of individual white spruce trees.

The resource budget model for mast seeding hypothesizes that soil nutrients proximately influence reproduction. Plants in high soil nutrient (particularly N) areas are predicted to have lower reproductive variability over time and higher mean reproduction. While often examined theoretically, there are relatively few empirical tests of this hypothesis. We quantified cone production of 110 individual white spruce (Picea glauca) trees over seven years and quantified plant-available soil macronutrients (N, Ca, K, Mg, P, S) in natural forest conditions across three years with different cone crop conditions. Each of these plant-available soil nutrients were correlated across years (rs = 0.55-0.89; all > 0.81 for total-N); spatially, total-N availability varied 366-fold across trees. Plant-available soil nutrients did not influence variability or mean annual reproduction, contrary to nutrient perturbation experiments. We examined within-year nutrient and cone-production relationships, and observed significant positive relationships between reproduction and plant-available soil nutrients only in a low-reproduction year preceding a mast event. Both during a mast event and the following year, when overall cone production was very high or very low, there were no relationships. Both external drivers (e.g., weather) and internal resource budgets likely influence soil nutrient-reproduction relationships. These results suggest that plant-available soil nutrients may not be a large factor influencing mast-seeding patterns among individuals in this species.

Formation of tussocks by sedges: effects of hydroperiod and nutrients.

Tussock formation is a global phenomenon that enhances microtopography and increases biodiversity by adding structure to ecological communities, but little is known about tussock development in relation to environmental factors. To further efforts to restore wetland microtopography and associated functions, we investigated Carex stricta tussock size in relation to elevation (a proxy for water depth) at a range of sites in southern Wisconsin, USA, and tested the effect of five hydroperiods and N+P addition (15 g N/m2 + 0.37 g P/m2) on tussock formation during a three-year mesocosm experiment. Wet meadows dominated by C. stricta averaged 4.9 tussocks/m2, with a mean volume of 1160 cm3 and height of 15 cm. Within sites, taller tussocks occurred at lower elevations, suggesting a structural adaptation to anoxic conditions. In our mesocosm experiment, C. stricta accelerated tussock formation when inundated, and it increased overall productivity with N + P addition. Within two growing seasons, continuous inundation (+18 cm) in the mesocosms led to tussocks that were nearly as tall as in our field survey (mean height in mesocosms, 10 +/- 1.3 cm; maximum, 17 cm). Plants grown with constant low water (-18 cm) only formed short mounds (mean height = 2 +/- 0.4 cm). After three growing seasons, the volume of the largest tussocks (3274 +/- 376 cm3, grown with +18 cm water depth and N + P addition) was 12 times that of the smallest (275 +/- 38 cm3, grown with -18 cm water depth and no N + P). Though tussock composition varied among hydroperiods, tussocks were predominantly organic (74-94% of dry mass) and composed of leaf bases (46-59%), fine roots (10-31%), and duff (5-13%). Only the plants subjected to high water levels produced the vertically oriented rhizomes and ascending shoot bases that were prevalent in field-collected tussocks. Under continuous or periodic inundation, tussocks achieved similar heights and accumulated similar levels of organic matter (range: 163-394 g C/m2), and we conclude that these hydroperiods can accelerate tussock formation. Thus, C. stricta has high utility for restoring wetland microtopography and associated functions, including carbon accumulation.

Repeated large-scale mechanical treatment of invasive Typha under increasing water levels promotes floating mat formation and wetland methane emissions.

Invasive species management typically aims to promote diversity and wildlife habitat, but little is known about how management techniques affect wetland carbon (C) dynamics. Since wetland C uptake is largely influenced by water levels and highly productive plants, the interplay of hydrologic extremes and invasive species is fundamental to understanding and managing these ecosystems. During a period of rapid water level rise in the Laurentian Great Lakes, we tested how mechanical treatment of invasive plant Typha × glauca shifts plant-mediated wetland C metrics. From 2015 to 2017, we implemented large-scale treatment plots (0.36-ha) of harvest (i.e., cut above water surface, removed biomass twice a season), crush (i.e., ran over biomass once mid-season with a tracked vehicle), and Typha-dominated controls. Treated Typha regrew with approximately half as much biomass as unmanipulated controls each year, and Typha production in control stands increased from 500 to 1500 g-dry mass m-2 yr-1 with rising water levels (~10 to 75 cm) across five years. Harvested stands had total in-situ methane (CH4) flux rates twice as high as in controls, and this increase was likely via transport through cut stems because crushing did not change total CH4 flux. In 2018, one year after final treatment implementation, crushed stands had greater surface water diffusive CH4 flux rates than controls (measured using dissolved gas in water), likely due to anaerobic decomposition of flattened biomass. Legacy effects of treatments were evident in 2019; floating Typha mats were present only in harvested and crushed stands, with higher frequency in deeper water and a positive correlation with surface water diffusive CH4 flux. Our study demonstrates that two mechanical treatments have differential effects on Typha structure and consequent wetland CH4 emissions, suggesting that C-based responses and multi-year monitoring in variable water conditions are necessary to accurately assess how management impacts ecological function.

Nitrogen enrichment stimulates wetland plant responses whereas salt amendments alter sediment microbial communities and biogeochemical responses.

Freshwater wetlands of the temperate north are exposed to a range of pollutants that may alter their function, including nitrogen (N)-rich agricultural and urban runoff, seawater intrusion, and road salt contamination, though it is largely unknown how these drivers of change interact with the vegetation to affect wetland carbon (C) fluxes and microbial communities. We implemented a full factorial mesocosm (378.5 L tanks) experiment investigating C-related responses to three common wetland plants of eastern North America (Phragmites australis, Spartina pectinata, Typha latifolia), and four water quality treatments (fresh water control, N, road salt, sea salt). During the 2017 growing season, we quantified carbon dioxide (CO2) and methane (CH4) fluxes, above- and below-ground biomass, root porosity, light penetration, pore water chemistry (NH4+, NO3-, SO4-2, Cl-, DOC), soil C mineralization, as well as sediment microbial communities via 16S rRNA gene sequencing. Relative to freshwater controls, N enrichment stimulated plant biomass, which in turn increased CO2 uptake and reduced light penetration, especially in Spartina stands. Root porosity was not affected by water quality, but was positively correlated with CH4 emissions, suggesting that plants can be important conduits for CH4 from anoxic sediment to the atmosphere. Sediment microbial composition was largely unaffected by N addition, whereas salt amendments induced structural shifts, reduced sediment community diversity, and reduced C mineralization rates, presumably due to osmotic stress. Methane emissions were suppressed by sea salt, but not road salt, providing evidence for the additional chemical control (SO4-2 availability) on this microbial-mediated process. Thus, N may have stimulated plant activity while salting treatments preferentially enriched specific microbial populations. Together our findings underpin the utility of combining plant and microbial responses, and highlight the need for more integrative studies to predict the consequences of a changing environment on freshwater wetlands.

Invasive species removal increases species and phylogenetic diversity of wetland plant communities.

Plant invasions result in biodiversity losses and altered ecological functions, though quantifying loss of multiple ecosystem functions presents a research challenge. Plant phylogenetic diversity correlates with a range of ecosystem functions and can be used as a proxy for ecosystem multifunctionality. Laurentian Great Lakes coastal wetlands are ideal systems for testing invasive species management effects because they support diverse biological communities, provide numerous ecosystem services, and are increasingly dominated by invasive macrophytes. Invasive cattails are among the most widespread and abundant of these taxa. We conducted a three-year study in two Great Lakes wetlands, testing the effects of a gradient of cattail removal intensities (mowing, harvest, complete biomass removal) within two vegetation zones (emergent marsh and wet meadow) on plant taxonomic and phylogenetic diversity. To evaluate native plant recovery potential, we paired this with a seed bank emergence study that quantified diversity metrics in each zone under experimentally manipulated hydroperiods. Pretreatment, we found that wetland zones had distinct plant community composition. Wet meadow seed banks had greater taxonomic and phylogenetic diversity than emergent marsh seed banks, and high-water treatments tended to inhibit diversity by reducing germination. Aboveground harvesting of cattails and their litter increased phylogenetic diversity and species richness in both zones, more than doubling richness compared to unmanipulated controls. In the wet meadow, harvesting shifted the community toward an early successional state, favoring seed bank germination from early seral species, whereas emergent marsh complete removal treatments shifted the community toward an aquatic condition, favoring floating-leaved plants. Removing cattails and their litter increased taxonomic and phylogenetic diversity across water levels, a key environmental gradient, thereby potentially increasing the multifunctionality of these ecosystems. Killing invasive wetland macrophytes but leaving their biomass in situ does not address their underlying mechanism of dominance and is less effective than more intensive treatments that also remove their litter.

Influence of nonnative earthworms on mycorrhizal colonization of sugar maple (Acer saccharum).

• Exotic earthworms can modify or eliminate surface organic (Oe/Oa) horizons in cold-temperate forest ecosystems and have profound effects on the forest soil environment, especially the rooting zone. • We examined the effects of earthworm colonization of northern hardwood forest soils on the abundance and morphology of mycorrhizal fungi associated with sugar maple ( Acer saccharum ). We compared mycorrhizal associations of areas of earthworm invasion with those of reference (no-worm) areas in Arnot Forest, central New York, USA. • The organic horizon in reference areas had higher mycorrhizal colonization rates and higher colonized root length than did surface layers in areas with active earthworm populations. Hyphal coils were more abundant and also formed a greater proportion of total fungal colonization in reference plots. Vesicles were more abundant and were a higher contribution to total colonization in earthworm plots, indicating a possible stress response to the presence of earthworms. • By affecting mycorrhizal colonization and morphology, earthworms may influence nutrient uptake capacity of dominant forest species. Our results suggest that a profound change in the mycorrhizal system will be one component of the potential ecosystem effects of invasion of new forest habitat by nonnative earthworms.

Intraepithelial gammadelta+ lymphocytes maintain the integrity of intestinal epithelial tight junctions in response to infection.

BACKGROUND & AIMS: Intestinal epithelial integrity and permeability is dependent on intercellular tight junction (TJ) complexes. How TJ integrity is regulated remains unclear, although phosphorylation and dephosphorylation of the integral membrane protein occludin is an important determinant of TJ formation and epithelial permeability. We have investigated the role intestinal intraepithelial lymphocytes (iIELs) play in regulating epithelial permeability in response to infection. METHODS: Recombinant strains of Toxoplasma gondii were used to assess intestinal epithelial barrier function and TJ integrity in mice with intact or depleted populations of iIELs. Alterations in epithelial permeability were correlated with TJ structure and the state of phosphorylation of occludin. iIEL in vivo reconstitution experiments were used to identify the iIELs required to maintain epithelial permeability and TJ integrity. RESULTS: In the absence of gammadelta+ iIELs, intestinal epithelial barrier function and the ability to restrict epithelial transmigration of Toxoplasma and the unrelated intracellular bacterial pathogen Salmonella typhimurium was severely compromised. Leaky epithelium in gammadelta+ iIEL-deficient mice was associated with the absence of phosphorylation of serine residues of occludin and lack of claudin 3 and zona occludens-1 proteins in TJ complexes. These deficiencies were attributable to the absence of a single subset of gammadelta T-cell receptor (TCR-Vgamma7+) iIELs that, after reconstituting gammadelta iIEL-deficient mice, restored epithelial barrier function and TJ complexes, resulting in increased resistance to infection. CONCLUSIONS: These findings identify a novel role for gammadelta+ iIELs in maintaining TJ integrity and epithelial barrier function that have implications for understanding the pathogenesis of intestinal inflammatory diseases associated with disruption of TJ complexes.