Salt Marsh Grass Spartina Alterniflora Research Articles

Self‐thinning and size‐dependent flowering of the grass Spartina alterniflora across space and time

Abstract Plants adjust their size and reproductive effort in response to numerous selection pressures and constraints. The self‐thinning law describes a well‐known trade‐off between size and density. Plants also trade‐off investment into growth vs. sexual reproduction, as described by life‐history theory. We build on past work on plant allometry and life history by examining both self‐thinning and size‐dependent reproduction in a single plant species, the saltmarsh grass Spartina alterniflora, across a wide range of settings: three landscape positions, two habitats and eight sites, across sixteen years. Plants in different landscape positions and years varied tremendously in size and shoot density. However, all this variation could be explained by a single allometric relationship consistent with the self‐thinning law, but with a lower slope. Flowering was size‐dependent, and the size at which plants had a 50% probability of flowering varied among habitat, sites and years. Plants that were stressed reproduced at a smaller size than plants that were growing under good conditions, and this pattern was consistent among habitat, sites and years. Finally, reproductive biomass and the proportion of shoots flowering increased with increasing vegetative size (plant height or shoot biomass). Combining these two patterns, S. alterniflora plants growing high density are small and reproduce at a smaller size than large plants growing at low density. Although there is tremendous spatial and temporal variation in S. alterniflora growth and reproductive patterns, all this variation can be understood as resulting from two simple allometric trade‐offs. Because saltmarsh plants often occur in monospecific stands, they may serve as simple, model systems for studies of plant life history. A free Plain Language Summary can be found within the Supporting Information of this article.

Nematode Responses to the Invasion of Exotic Spartina in Mangrove Wetlands in Southern China

Investigations into nematode density and species assemblages have been conducted in different types of mangroves worldwide, but these studies have typically been limited to one type of plant or tree species. The invasive salt marsh grass Spartina alterniflora has successively invaded native mangroves along the southern coasts of China during the preceding two decades. However, few meiofauna studies on the impacts of S. alterniflora have been conducted, and the consequences of this invasion on ecosystem composition and function remain unclear. The hypothesis of this study was that the spatial and seasonal distribution of nematode assemblages vary significantly among three native mangrove habitats (Kandelia obovata, Aegiceras corniculatum, and Avicennia marina) and between these habitats and a fourth habitat that was colonized by S. alterniflora, in Zhangjiang Estuary, China. Our results demonstrated that different species dominated in different habitats seasonally. Highly significant differences in density, number of species, diversity index, and maturity index were present among the four habitats. ANOSIM results revealed that there were significant differences in nematode assemblages among the four habitats and seasons, with the S. alterniflora habitat exhibiting the lowest mean values of number of species, Shannon-Wiener diversity index, richness index, and maturity index in the four seasons. This suggests that the presence of S. alterniflora disrupted nematode assemblages.

Grazing Scar Characteristics Impact Degree of Fungal Facilitation in Spartina alterniflora Leaves in a South American Salt Marsh

Grazing scars of burrowing crabs and Hemiptera insects were simulated on leaves of the salt marsh grass Spartina alterniflora. Simulations of crab feeding generated two-fold higher fungal (ergosterol) content in leaves in comparison to that generated by insect scar simulations (1.26 ±0.55 and 0.57 ±0.25 µg per cm², respectively). This study provided evidence that herbivory could facilitate microbial infection by fungi in dominant South American salt marsh plants and indicated that specific feeding mechanisms used by different herbivores might differentially impact the strength of this interaction.

Delimiting the coastal geographic background to predict potential distribution of Spartina alterniflora

Ecological niche modeling is an important tool in studying biological invasion; however, the geographic background on niche model transferability received scant attention. The salt marsh grass Spartina alterniflora, natively distributed along the eastern coasts of the Americas, is considered a global invasive species. In this study, we first compared the climate space among geographically separated populations. The classic niche model approaches involving the calibration of native range climate envelop of S. alterniflora and transferring worldwide were then used to predict potential invasion. Niche models based on two geographic backgrounds were compared, namely a large squared area delimited by a bounding box containing all known occurrences, which is usually used in former studies and a small coastal area defined as the geographic space available to the species. Both area-based models showed good performance in native range predictions, however, when models were transferred, niche model calibrated on the coastal area showed higher predictability in capturing the introduced occurrences. Given the potential substantial effect of geographic background on niche model transferability, caution is warranted when interpreting low-niche model transferability with niche differentiation and when predicting other (coastal) species’ invasion.

Hydrologic variability in a salt marsh: Assessing the links between drought and acute marsh dieback

It has been hypothesized that acute marsh dieback (AMD) observed along the Gulf Coast and South Atlantic Bight in the early 2000s was the result of drought-induced changes to porewater and sediment chemistry through hypersalinity or through mobilization of metals and acidification associated with redox changes. The impact of drought on coastal wetlands remains unclear because the hydrology of these wetlands is strongly influenced by regular tidal inundation. In order to test the links between hydrologic variability and changes to marsh groundwater conditions that may be stressful to the salt marsh grass Spartina alterniflora, we installed piezometers and passive diffusion samplers in a salt marsh island at North Inlet, South Carolina, where AMD was observed in fall 2001. Significant variations in tidal inundation, rainfall, evapotranspiration, groundwater dynamics, and porewater chemistry were observed. The island was typically inundated twice daily, but there were occasional 19–21 h periods in winter and spring when the marsh was not inundated and a singular event when the marsh was not inundated for three days (March 2008). Enhanced exposure resulted in seasonal redox chemistry changes, as indicated by changes in the ratio of ferrous iron [Fe(II)] to total iron [Fe(II) + Fe(III)], but our observations do not support redox and pH changes as the cause of AMD at this site. Porewater salinity varied from 14 to 40 in the upper 1 m of the marsh. Salinity was most variable near the surface and increased with depth, reflecting root zone transpiration and downward movement of porewater through the marsh mud into the underlying confined sand aquifer. Pearson Correlation tests among porewater constituents and hydrologic parameters indicated significant associations between porewater salinity, tidal inundation, rainfall, and ET, and additional associations between porewater iron concentration, speciation, and tidal inundation. Linear regression model estimates of porewater salinity for 2001–2002 did not indicate the development of hypersalinity during that period. However, these estimates did predict a dramatic increase in salinity that coincided with the beginning of drought conditions just prior to the observation of AMD, suggesting this as a cause for AMD at this site. Drought is predicted to increase over the next century; damage caused by potential increases in the frequency of drought-related AMD may limit the ability of intertidal salt marshes to accommodate sea level rise.

SURVIVAL AND GROWTH OF THE DOMINANT SALT MARSH GRASS SPARTINA ALTERNIFLORA IN AN OIL INDUSTRY SALINE WASTEWATER

Saline oil produced water (PW) is the largest wastewater stream in the oil exploration and production processes. Although eventual disposal of PW into shallow coastal waters occurs nearby coastal wetlands, no studies regarding its toxicity to higher plants were found in our literature review. To fill this knowledge gap and evaluate the potential use of this halophyte for PW phytoremediation the salt marsh grass Spartina alterniflora was grown in five PW concentrations and no PW treatment control for seven weeks. The oil & grease, NaCl, and ammonium (N-NH4+) concentrations in the PW were 120 mg L−1, 30 g L−1, and 381 mg L−1, respectively. Plants grown in 30% PW and 10% PW achieved survival rates (75%) significantly higher than plants grown in 100% PW (35% survival). LT50 of S. alterniflora to raw PW with 120 mg L−1 of oil & grease (100% PW) was estimated at 30 days. Root and sprout biomass were significantly stimulated by PW; plants grown in 10% to 50% PW concentrations were 70–300% more productive than those in control, 80% PW and 100% PW, respectively. No significant inhibitory effects on survival or growth were detected for concentrations of PW less than 80% when compared to control. Our results pointed out that S. alterniflora grows in saline oil PW and its potential use to phytoremediate this effluent should be evaluated.

Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient

An invasive variety of Phragmites australis (Poaceae, common reed), the M haplotype, has been implicated in the spread of this species into North American salt marshes that are normally dominated by the salt marsh grass Spartina alterniflora (Poaceae, smooth cordgrass). In some European marshes, on the other hand, Spartina spp. derived from S. alterniflora have spread into brackish P. australis marshes. In both cases, the non-native grass is thought to degrade the habitat value of the marsh for wildlife, and it is important to understand the physiological processes that lead to these species replacements. We compared the growth, salt tolerance, and osmotic adjustment of M haplotype P. australis and S. alterniflora along a salinity gradient in greenhouse experiments. Spartina alterniflora produced new biomass up to 0.6 M NaCl, whereas P. australis did not grow well above 0.2 M NaCl. The greater salt tolerance of S. alterniflora compared with P. australis was due to its ability to use Na(+) for osmotic adjustment in the shoots. On the other hand, at low salinities P. australis produced more shoots per gram of rhizome tissue than did S. alterniflora. This study illustrates how ecophysiological differences can shift the competitive advantage from one species to another along a stress gradient. Phragmites australis is spreading into North American coastal marshes that are experiencing reduced salinities, while Spartina spp. are spreading into northern European brackish marshes that are experiencing increased salinities as land use patterns change on the two continents.

Aboveground and belowground productivity ofSpartina alterniflora (Smooth Cordgrass) in natural and created Louisiana salt marshes

In Louisiana, salt marshes are being created in an effort to offset the large loss of such habitat that has occurred over the last 50 yr. Primary productivity is an important function and indicator of success for salt marsh creation and restoration projects. The aim of this study was to determine whether the aboveground and belowground productivity of the dominant salt marsh grassSpartina alterniflora in created marshes in southwest Louisiana began to approximate productivity levels in natural marshes, over time. Net annual aboveground primary productivity (NAPP) was measured by a harvest technique, while the ingrowth core method was used to estimate net annual belowground primary productivity (NBPP). NAPP levels were similar to those found in other, Louisiana salt marshes, while NBPP levels were similar to or higher than the reported range forS. alterniflora studied along the Atlantic and Gulf of Mexico coasts. NAPP tended to decrease as the created marshes aged, but the levels in the oldest, 19 year old, created marsh were still well above values measured in the, natural marshes. It was estimated that it would take 35 yr after marsh creation for NAPP in the created marshes to become equivalent to that in natural marshes. NBPP in the created marshes became equivalent to levels found in the natural marshes after 6–8 yr, but then belowground production increased with marsh age, reaching an asymptote that surpassed natural marsh levels. Equivalency in primary productivity has not been reached in these marshes. Elevation also affected productivity, as higher elevational sites with greater topographic heterogeneity had significantly lower aboveground and belowground biomass levels than those with elevations closer to mean sea level. This underscores the need to construct marshes so that their mean elevation and degree of topographic heterogeneity are similar to natural marshes.

Dimethylsulfoniopropionate biosynthesis in Spartina alterniflora1. Evidence that S-methylmethionine and dimethylsulfoniopropylamine are intermediates.

The osmoprotectant 3-dimethylsulfoniopropionate (DMSP) occurs in Gramineae and Compositae, but its synthesis has been studied only in the latter. The DMSP synthesis pathway was therefore investigated in the salt marsh grass Spartina alterniflora Loisel. Leaf tissue metabolized supplied [35S]methionine (Met) to S-methyl-l-Met (SMM), 3-dimethylsulfoniopropylamine (DMSP-amine), and DMSP. The 35S-labeling kinetics of SMM and DMSP-amine indicated that they were intermediates and, consistent with this, the dimethylsulfonium moiety of SMM was shown by stable isotope labeling to be incorporated as a unit into DMSP. The identity of DMSP-amine, a novel natural product, was confirmed by both chemical and mass-spectral methods. S. alterniflora readily converted supplied [35S]SMM to DMSP-amine and DMSP, and also readily converted supplied [35S]DMSP-amine to DMSP; grasses that lack DMSP did neither. A small amount of label was detected in 3-dimethylsulfoniopropionaldehyde (DMSP-ald) when [35S]SMM or [35S]DMSP-amine was given. These results are consistent with the operation of the pathway Met --> SMM --> DMSP-amine --> DMSP-ald --> DMSP, which differs from that found in Compositae by the presence of a free DMSP-amine intermediate. This dissimilarity suggests that DMSP synthesis evolved independently in Gramineae and Compositae.

The Biogeochemistry and Trace Metals Distribution of Mangrove Rhizospheres

Soils and porewater of the rhizosphere of mangrove trees, Rhizophora mangle L. and Avicennia schaueriana Stapf & Leech., of the salt marsh grass Spartina alterniflora Loisel, and of unvegetated mud flats, were analyzed for pH, salinity, platinum electrode redox potential, organic matter content, sulfide concentrations, and the total and exchangeable concentrations of trace metals (Fe, Zn, Cu, Pb and Cd). The study was conducted in a fringe mangrove forest in Sepetiba Bay, Rio de Janeiro, Brazil. The rhizospheres differed in their biogeochemistry. Mud flats and Rhizophora soils were very reducing, with highest concentrations of sulfide. Avicennia soils showed the highest variability of the variables measured, with the rhizosphere changing from oxic to anoxic conditions. Spartina soils, on the other hand, were generally oxic, with very low sulfide concentration. The distribution of trace metals in these soils varied with the major physical and chemical characteristics. Mud flat soils presented the highest total trace metal concentrations followed by mangrove soils and Spartina soils. However, exchangeable trace metals were similar among the different soils with the exception of Avicennia soils, which due to their characteristic instability of redox conditions, presented much higher exchangeable trace metals concentrations.

Effect of salinity on the critical nitrogen concentration of Spartina alterniflora Loisel

Nitrogen was withheld from the salt marsh grass Spartina alterniflora Loisel., in order to determine the effect of salinity (sea salts) on critical tissue nitrogen concentrations (defined here as the minimum tissue concentration required to sustain biomass accumulation). The critical nitrogen concentration per kilogram dry weight of above-ground tissue increased non-linearly from a mean of 8.2 g kg −1 at 5 g l −1 and 20 g l −1 salinity to 13.6 g kg −1 and 22.9 g kg −1 at salinities of 40 g l −1 and 50 g l −1, respectively. Below-ground tissue nitrogen concentrations averaged 62% of the above-ground values irrespective of salinity treatment. These results suggest that the critical nitrogen concentration is a function of salinity and indicate that the internal nitrogen supply required in support of growth increases with salinity. Above-ground tissue nitrogen concentrations reported in the literature and the relationship between salinity and critical nitrogen concentration observed in this study were used to evaluate the nitrogen status of S. alterniflora over a wide range of geographical locations. Comparisons suggest that both short and tall forms of S. alterniflora are nitrogen limited in the majority of marshes along the Gulf and Atlantic Coasts of the US.

Tight coupling of root-associated nitrogen fixation and plant photosynthesis in the salt marsh grass Spartina alterniflora and carbon dioxide enhancement of nitrogenase activity.

The coupling of root-associated nitrogen fixation and plant photosynthesis was examined in the salt marsh grass Spartina alterniflora. In both field experiments and hydroponic assay chambers, nitrogen fixation associated with the roots was rapidly enhanced by stimulating plant photosynthesis. A kinetic analysis of acetylene reduction activity (ARA) showed that a five-to sixfold stimulation occurred within 10 to 60 min after the plant leaves were exposed to light or increased CO2 concentrations (with the light held constant). In field experiments, CO2 enrichment increased plant-associated ARA by 27%. Further evidence of the dependence of ARA on plant photosynthate was obtained when activity in excised roots was shown to decrease after young greenhouse plants were placed in the dark. Seasonal variation in the ARA of excised plant roots from field cores appears to be related to the annual cycle of net photosynthesis in S. alterniflora.

Lignocellulose and lignin in the salt marsh grass Spartina alterniflora: initial concentrations and short-term, post-depositional changes in detrital matter

Lignocellulose was found to comprise the bulk of various anatomical structures of the salt marsh grass Spartina alterniflora, as well as of detritus derived from this plant; although concentrations of both lignin and lignocellulose varied with plant height-form, age, and anatomical structure. Changes in the relative concentrations of lignin and polysaccharide in lignocellulose due to long-term degradation by natural marsh microflora were determined using standard gravimetric assay procedures and a new procedure utilizing specifically-radiolabelled 14C-(cellulose)-lignocellulose and 14C-(lignin)-lignocellulose prepared from S. alterniflora. Results obtained with the two methods agreed, thus validating the new procedures. Mineralization of the cellulose moiety was more rapid than mineralization of the lignin moiety resulting in relative enrichment of S. alterniflora detritus in lignin. Rates of mineralization of both moieties decreased over time such that extent of mineralization was a function of the logarithm of aging time.

Effects of salinity and illumination on photosynthesis and water balance of Spartina alterniflora Loisel.

Plants of the salt marsh grass Spartina alterniflora Loisel were collected from North Carolina and grown under controlled nutrient, temperature, and photoperiod conditions. Plants were grown at two different illumination levels; substrate salinity was varied, and leaf photosynthesis, transpiration, total chlorophyll, leaf xylem pressure, and specific leaf weight were measured. Conditions were controlled so that gaseous and liquid phase resistances to CO2 diffusion could be calculated. Growth at low illumination and high salinity (30 ppt) resulted in a 50% reduction in photosynthesis. The reduction in photosynthesis of plants grown at low illumination was correlated with an increase in gaseous resistance. Photosynthetic rates of plants grown at high salinity and high illumination were reduced only slightly compared to rates of plants grown, in 10 ppt and Hoagland's solution. Both high salinity and high illumination were correlated with increases in specific leaf weight. Chlorophyll data indicate that specific leaf weight differences were the result of increases in leaf thickness. It is therefore hypothesized that photosynthetic response can be strongly influenced by salinity-induced changes in leaf structure. Similarities in photosynthetic rate on an area basis at high, illumination were apparently the result, of increases in leaf thickness at high salinity. Photosynthetic rates were generally quite high, even at salinities close to open ocean water, and it is concluded that salinity rarely limits photosynthesis in S. alterniflora.

SEED GERMINATION RESPONSE AND EVIDENCE FOR HEIGHT ECOPHENES IN SPARTINA ALTERNIFLORA FROM NORTH CAROLINA

A study was conducted to determine germination response to temperature and salinity and seedling response to salinity by three height forms of the salt marsh grass Spartina alterniflora Loisel. Germination tests showed that seeds cannot withstand drying at moderate temperature, as viability was lost within 40 days in seeds stored dry at 72 F. Cold storage at 43 F is adequate to prevent desiccation up to 40 days, but after 8 months viability is lost. Viability is retained at least 8 months when seeds are stored in sea water at 43 F. Germination response was good in a 65–95 F alternating diurnal thermoperiod but was poor in a 72 F constant thermoperiod. Germination response to salinity was an inverse curvilinear relationship with germination inhibition at high salinities apparently due to osmotic effects. The maximum tolerance limit for germination lies between 6 and 8 % NaCl. Seeds from short, medium, and tall plants responded similarly in storage and temperature studies. However, in salinity studies, seeds of the Ocracoke Island short form and the Oak Island tall form performed best. A logarithmic curve best described seedling growth response to various NaCl levels. Growth response as measured by seedling dry weight was best in 0.5 % NaCl solution. Seedlings grew taller in both 0.5 and 1.0 % NaCl than in 0 % NaCl. No significant difference in seedling growth response due to height form of the parent plant was detected. Thus, on the basis of germination and seedling responses, the height forms of S. alterniflora in North Carolina salt marshes are best described as ecophenes.