Effect of the Spartina alterniflora Root-Rhizome System on Salt Marsh Soil Denitrifying Bacteria

Nitrous oxide (N₂O) reductase activity was used as an index of the denitrification potential in salt marsh soils. In a short Spartina alterniflora marsh, the seasonal distribution of N₂O reductase activity indicated a causal relationship between S. alterniflora root-rhizome production and the denitrification potential of the soil system. The relationship was not discerned in samples from a tall S. alterniflora marsh. To further examine the in situ plant-denitrifier interaction in the short S. alterniflora marsh, plots with and without living S. alterniflora were established and analyzed for N₂O reductase activity 5 and 18 months later. In the plots without living Spartina there was a significant reduction in the soil denitrification potential after 18 months, indicating that in the SS marsh the denitrifiers are tightly coupled to the seasonal production of below-ground Spartina macroorganic matter. In plots with intact Spartina, the soil denitrification potential was not altered by NH₄NO₃ or glucose enrichment. However, in plots without living Spartina, there were significant changes in soil N₂O reductase activity, thus indicating that the plants can serve as a “buffer” against this form of pulse perturbation.     

Role of the Salt Marsh Grass Spartina alterniflora in the Response of Soil-Denitrifying Bacteria to Glucose Enrichment

Long-term incubations of salt marsh soil systems in the presence of glucose resulted in a decrease in the soils' denitrification potential. Addition of nitrate or the presence of living Spartina alterniflora reversed this effect, indicating that Spartina, through the establishment of an oxidized rhizosphere where nitrification can occur, enables the denitrifying bacteria to adequately compete with the less energetically efficient components of the anaerobic soil microbial community.     

Populations of Methanogenic Bacteria in a Georgia Salt Marsh

Methanogens represented about 0.5% of the total bacteria in sediments from a Georgia salt marsh in which Spartina alterniflora is the predominant vegetation. The population of methanogens was composed of at least two groups of nearly equal size. One group was represented by cocci which were able to utilize trimethylamine and were unable to use H₂ or acetate. The second group was composed of two subgroups which were able to utilize H₂ but were unable to use trimethylamine or acetate. The more common subgroup included rod- or plate-shaped methanogens which could utilize isopropanol in addition to H₂ and formate. The second subgroup included Methanococcus maripaludis, which utilized only H₂ and formate. Other groups of methanogens were also present, including Methanosarcina sp. which utilized acetate, H₂, and methylamines. In addition to the overall variability in the types of methanogens, the numbers of methanogens in sediments also exhibited significant spatial variability both within and between tall- and short-Spartina zones.     

Comparison of Nitrogen Fixation Activity in Tall and Short Spartina alterniflora Salt Marsh Soils

A comparison of the N₂ fixers in the tall Spartina alterniflora and short S. alterniflora marsh soils was investigated. Zero-order kinetics and first-order kinetics of acetylene reduction were used to describe the activity of the N₂ fixers in marsh soil slurries. It was found that the V_max values were approximately 10 times greater for the N₂ fixers in the tall Spartina than in the short Spartina marsh when raffinose was used as the energy source. In addition, the (K_s + S_n) values were approximately 4 to 15 times lower for the N₂ fixers in the tall Spartina than in short Spartina marsh. First-order kinetics of nitrogen fixation for several substrates indicate that the N₂ fixers in the tall Spartina marsh were two to seven times more active than those in the short Spartina marsh. Ammonium chloride (25 μg/ml) did not inhibit nitrogen fixation in the tall Spartina marsh, but there was a 50% inhibition in nitrogen fixation in the short Spartina marsh. On the other hand, sodium nitrate inhibited nitrogen fixation almost 100% at 25 μg/ml in both soil environments. Amino nitrogen (25 to 100 μg/ml) had little or no effect on nitrogen fixation. The results indicate that the N₂ fixers in the tall Spartina marsh were physiologically more responsive to nutrient addition than those in the short Spartina marsh. This difference in the two populations may be related to the difference in daily tidal influence in the respective areas and thus provide another explanation for the enhanced S. alterniflora production in the creek bank soil system.     

沼泽湿地互花米草植物体传输与排放甲烷特征

于各测定月的小潮日,使用悬管装置采集气样并结合气相色谱测定,对闽江河口互花米草（Spartina alterniflora）沼泽湿地互花米草植株的甲烷传输量及其主要释放部位进行了研究,另又采用静态箱法-气相色谱法测定了互花米草沼泽湿地甲烷的排放通量,以此分析互花米草植物体甲烷传输对互花米草沼泽湿地甲烷排放通量的贡献率,最后测定了互花米草植株髓腔内的甲烷浓度。结果表明,不同生长阶段的互花米草植株甲烷传输量明显不同,快速生长阶段的互花米草植株甲烷传输量最高;互花米草植株甲烷传输对互花米草沼泽湿地甲烷排放通量的贡献率介于9%-94%之间;互花米草植物体传输甲烷的主要释放部位是距地面0-20cm处,该部位对植物甲烷传输量的贡献率为50%（平均值）;髓腔内的甲烷浓度远高于大气中的甲烷浓度,且自下而上呈递减趋势。     

Gas Transport in the Marsh Grass, Spartina alterniflora

Spartina plants have continuous gas spaces from the leaves tothe tips of the roots. Oxygen values in the roots are as lowas 3 per cent and increase toward the stem. Carbon dioxide valuesare highest in the rhizome and decrease up the stem and towardthe root tips. Oxygen and carbon dioxide moved through the plantsat equal rates for equal gradients, and these rates agree withmeasurements made on plants in the marsh. Calculated oxygenand carbon dioxide fluxes for the observed gradients in theobserved gas spaces agreed with measured fluxes. We concludethat gases move in and out of Spartina roots by diffusion throughuninterrupted gas spaces within the plant.     

Survivorship and Spatial Development of Spartina alterniflora Loisel. (Gramineae) Seedlings in a New England Salt Marsh

Patterns of survival and spatial arrangement of tillers of Spartinaalterniflora were examined in natural and in artificially producedbare areas, and in pure stands of adult S. alterniflora in aNew England salt marsh. Seedling growth and survival were highin naturally occurring bare patches and in artificial patches,whether created by continual clipping of adult plants to groundlevel throughout the growing season or by providing bare substrateafter removal of adult plants. Seedling growth and survivorshipincreased with increasing size of bare area. S. alternifloraseedlings were also common in areas dominated by adult plants,but no seedlings survived a whole first growing season underthe mature canopy, probably because of competition from adultplants. In large bare areas, S. alterniflora seedlings grew non-directionally,reaching heights of 0.5 m, and producing as many as 36 tillersin one growing season. Examination of leaf area ratios suggestedthat the production of tillers increased photosynthetic capacityin seedlings with several tillers in contrast to seedlings withouttillers. Thus, seedlings can apparently tiller and colonizefree space radially without a loss of photosynthetic capacity. These results suggest that while seedling success is generallylimited by adult plants in monocultures of S. alterniflora,in disturbed spaces seedling success is high and results ina rapid non-directional proliferation of sexually generatedclones that ultimately preclude the successful invasion of futureseedlings. Clones, directional growth, leaf area ratio, salt marsh cordgrass, seedling survivorship, Spartina alterniflora Loisel     

Seasonal Variability of Diazotroph Assemblages Associated with the Rhizosphere of the Salt Marsh Cordgrass, Spartina alterniflora

Nitrogen fixation is the primary N source in the highly productive but N-limited North Inlet, SC, USA salt marsh system. The diverse assemblages of nitrogen-fixing (diazotrophic) bacteria associated with the rhizospheres of the short and tall growth forms of Spartina alterniflora were analyzed at two sites, Crab Haul Creek and Goat Island, which are in different tidal creek drainage systems in this marsh. The sites differed in proximity to the main channel for tidal intrusion and in several edaphic parameters. We hypothesized that either the differing abiotic environmental regimes of the two sites or the variation due to seasonal effects result in differences in the diazotroph assemblage. Rhizosphere samples were collected seasonally during 1999 and 2000. DNA was purified and nifH amplified for denaturing gradient gel electrophoresis (DGGE) analysis of diazotroph assemblage composition. Principal components analysis was used to analyze the binary DGGE band position data. Season strongly influenced assemblage composition and biplots were used to identify bands that significantly affected the seasonal and site-specific clustering. The types of organisms that were most responsive to seasonal or site variability were identified on the basis of DGGE band sequences. Seasonally responsive members of the anaerobic diazotrophs were detected during the winter and postsenescence conditions and may have been responsible for elevated pore water sulfide concentrations. Sequences from a diverse assemblage of Gammaproteobacteria were predominant during growth periods of S. alterniflora. Abiotic environmental parameters strongly influenced both the S. alterniflora and the diazotrophic bacterial assemblages associated with this keystone salt marsh plant species.     

Microdiversity of Culturable Diazotrophs from the Rhizoplanes of the Salt Marsh Grasses Spartina alterniflora and Juncus roemerianus

Abstract Salt marshes dominated by Spartina alterniflora (smooth cordgrass) are among the most productive ecosystems known, despite nitrogen limitation. Rhizoplane/rhizosphere diazotrophy (nitrogen fixation) serves as a significant source of combined nitrogen in these systems. Several recent studies have demonstrated remarkable physiological and phylogenetic macro- and microdiversity within this important functional group of organisms. However, the ecological significance of this diversity is presently unknown. The physiological characteristics of the culturable, oxygen-utilizing fraction of the rhizoplane diazotroph assemblages from Spartina alterniflora and from another salt marsh grass, the black needle rush Juncus roemerianus, were examined in combination with an assessment of the phylogenetic relatedness by whole genome DNA–DNA hybridization. Analysis of substrate utilization data permitted quantitative evaluation of fully cross-hybridizing strain groups and physiological clusters. Phylogenetically related strains, defined by DNA homology ≥90% relative to the positive control, displayed extensive physiological diversity. Seven bootstrap-supported physiological clusters, composed largely of phylogenetically dissimilar strains, showed similar utilization patterns for at least one class of ecologically relevant substrates (carbohydrates, carboxylic acids, or amino acids). These diazotrophs appear to be physiologically adapted for utilization of specific substrates or classes of substrates, lending support to diazotrophic functional redundancy. Microenvironmental heterogeneity is credited for promoting this diversity by selecting for physiologically specialized diazotroph populations to occupy defined niches in situ. One outcome of this physiological diversity is maintenance of a crucial environmental function (nitrogen fixation) over a broad range of environmental conditions. Received: 15 October 1999; Accepted: 28 December 1999; Online Publication: 25 April 2000     

Populations of Methane-Producing Bacteria and In Vitro Methanogenesis in Salt Marsh and Estuarine Sediments

Most probable numbers (MPNs) of methanogens in various salt marsh and estuarine sediments were determined with an anaerobic, habitat-simulating culture medium with 80% H₂ plus 20% CO₂ as substrate. Average MPNs for the short Spartina (SS) marsh sediments of Sapelo Island, Ga., were maximal at the 5- to 7-cm depth (1.2 × 10⁷/g of dry sediment). Populations decreased to approximately 880/g of dry sediment at the 34- to 36-cm depth. There was no significant difference between summer and winter populations. In tall Spartina (TS) marsh sediments, average populations were maximal (1.2 × 10⁶/g of dry sediment) in the upper 0- to 2-cm zone; populations from the 5- to 36-cm zones were similar (average of 9 × 10⁴/g of dry sediment). Methanogenic populations for TS sediments of James Island Creek marsh, Charleston, S.C., were similar (average of 3 × 10⁶/g of dry sediment) for all depths tested (0 to 22 cm), which was comparable to the trend observed for TS sediments at Sapelo Island, Ga. Sediment grab samples collected along a transect of James Island Creek and its adjacent Spartina marsh had MPNs that were approximately 20 times greater for the region of Spartina growth (average of 10⁶/g of dry sediment) compared with the channel (approximately 5 × 10⁴ methanogens per g of dry sediment). A similar trend was found at Pawley's Island marsh, S.C., but populations were approximately one order of magnitude lower. In vitro rates of methanogenesis with SS sediments incubated under 80% H₂-20% CO₂ showed that the 5- to 7-cm region exhibited maximal activity (51 nmol of CH₄ g⁻¹ h⁻¹), which was greater than rates for sediments above and below this depth. SS sediment samples (5 to 7 cm) incubated under 100% N₂ and supplemented with formate exhibited rates of methanogenesis similar to those generated by samples under 80% H₂-20% CO₂. Replacing the N₂ atmosphere with H₂ resulted in an eightfold decrease in the rate of methanogenesis. In vitro methanogenic activity by TS salt marsh sediments, incubated under 80% H₂-20% CO₂, was similar for all depths tested (0 to 22 cm). TS sediment samples (0 to 7 cm) supplemented with formate and incubated under 100% N₂ had greater rates of methanogenesis compared with unsupplemented samples.     

Changes in Ecosystem Nitrogen and Carbon Allocation with Black Mangrove (Avicennia germinans) Encroachment into Spartina alterniflora Salt Marsh

Ecosystems - Increases in temperature are expected to facilitate encroachment of tropical mangrove forests into temperate salt marshes, yet the effects on ecosystem services are understudied. Our...     

Incorporation of Exotic Spartina alterniflora into Diet of Deposit-Feeding Snails in the Yangtze River Estuary Salt Marsh: Stable Isotope and Fatty Acid Analyses

The response of deposit-feeding animals to plant invasions is still unclear, because their food sources are often difficult to identify. We examined the effect of the exotic plant species, Spartina alterniflora, on the food source composition of two dominant snail species, Assiminea latericea and Cerithidea largillierti, in the Yangtze River estuary salt marsh using a combination of stable isotope and fatty acid analyses. We collected the snails and their potential food materials (sediment organic matter, particulate organic matter, and plant material) in S. alterniflora and native plant Phragmites australis marshes and then determined the composition of food sources of snails based on fatty acid markers and stable isotope composition. Our results indicated that A. latericea and C. largillierti are deposit feeders grazing on sedimentary particles originating from diatoms, bacteria, and vascular plants. Invasive S. alterniflora did not result in a change in the relative contribution of microalgae, bacteria, and vascular plants to the food source of the snails. Spartina alterniflora was confirmed to be assimilated by both snail species. The higher assimilation of S. alterniflora by A. latericea compared with C. largillierti is probably related to the greater ability of A. latericea to assimilate plant materials from detritus, as evidenced by fatty acid composition. Overall, S. alterniflora can be incorporated into the food web of the estuarine salt marsh by the dominant snail species with generalist-feeding habits.     

Molecular Phylogenetic and Biogeochemical Studies of Sulfate-Reducing Bacteria in the Rhizosphere of Spartina alterniflora

The population composition and biogeochemistry of sulfate-reducing bacteria (SRB) in the rhizosphere of the marsh grass Spartina alterniflora was investigated over two growing seasons by molecular probing, enumerations of culturable SRB, and measurements of SO₄²⁻ reduction rates and geochemical parameters. SO₄²⁻ reduction was rapid in marsh sediments with rates up to 3.5 μmol ml⁻¹ day⁻¹. Rates increased greatly when plant growth began in April and decreased again when plants flowered in late July. Results with nucleic acid probes revealed that SRB rRNA accounted for up to 43% of the rRNA from members of the domain Bacteria in marsh sediments, with the highest percentages occurring in bacteria physically associated with root surfaces. The relative abundance (RA) of SRB rRNA in whole-sediment samples compared to that of Bacteria rRNA did not vary greatly throughout the year, despite large temporal changes in SO₄²⁻ reduction activity. However, the RA of root-associated SRB did increase from <10 to >30% when plants were actively growing. rRNA from members of the family Desulfobacteriaceae comprised the majority of the SRB rRNA at 3 to 34% of Bacteria rRNA, with Desulfobulbus spp. accounting for 1 to 16%. The RA of Desulfovibrio rRNA generally comprised from <1 to 3% of the Bacteria rRNA. The highest Desulfobacteriaceae RA in whole sediments was 26% and was found in the deepest sediment samples (6 to 8 cm). Culturable SRB abundance, determined by most-probable-number analyses, was high at >10⁷ ml⁻¹. Ethanol utilizers were most abundant, followed by acetate utilizers. The high numbers of culturable SRB and the high RA of SRB rRNA compared to that of Bacteria rRNA may be due to the release of SRB substrates in plant root exudates, creating a microbial food web that circumvents fermentation.     

End Products of Anaerobic Chitin Degradation by Salt Marsh Bacteria as Substrates for Dissimilatory Sulfate Reduction and Methanogenesis

The anaerobic pathway of chitin decomposition by chitinoclastic bacteria was examined with an emphasis on end product coupling to other salt marsh bacteria. Actively growing chitinoclastic bacterial isolates produced primarily acetate, H₂, and CO₂ in broth culture. No sulfate-reducing or methanogenic isolates grew on chitin as sole carbon source or produced any measurable degradation products. Mixed cultures of chitin degraders with sulfate reducers resulted in positive sulfide production. Mixed cultures of chitin-degrading isolates with methanogens resulted in the production of CH₄ with reductions in headspace CO₂ and H₂. The combination of all three metabolic types resulted in the simultaneous production of methane and sulfide, with more methane being produced in mixed cultures containing CO₂-reducing methanogens and acetoclastic sulfate reducers because of less interspecific H₂ competition.     

Naked Amoebas and Bacteria in an Oil-Impacted Salt Marsh Community

Populations of soil amoebas were monitored in two salt marshes in Staten Island, NY for 2 years. One site, Gulfport Reach on the Arthur Kill, has been highly impacted by numerous oil spills. In particular, in 1990 a massive no. 2 fuel oil spill from a ruptured pipe flooded the area; its sediments had total petroleum hydrocarbon (TPH) concentrations in the range 800-46,000 ppm. A reference site 11 km away, on the Atlantic coast, had low TPH levels. Amoeba population densities were in general higher in the impacted sediments. In laboratory microcosm experiments, sediment samples from unimpacted sites were treated with added fresh (unweathered) hydrocarbons (no. 2 fuel oil) and cultured; these also yielded higher amoeba numbers than untreated controls. Four distinct amoeba morphotypes were monitored. Changes in population levels of total amoebas were correlated in the two sites, particularly for morphotype 2 (r = 0.83). The ratios of total amoebas to total bacterial numbers were also correlated (r = 0.85) between the sites. This suggests the amoebas may function as generalists, and that their trophic relation to bacterial prey is not much affected by the presence of petroleum hydrocarbons, but rather may reflect regional parameters such as ambient temperature or other physical factors.     

Restoration of Arthropod Assemblages in a Spartina Salt Marsh following Removal of the Invasive Plant Phragmites australis

Invasive plants are one of the most serious threats to native species assemblages and have been responsible for the degradation of natural habitats worldwide. As a result, removal of invasive species and reestablishment of natural vegetation have been attempted in order to restore biodiversity and ecosystem function. This study examined how native arthropod assemblages, an abundant and functionally important group of organisms in many ecosystems, are affected by the incursion of the invasive wetland plant Phragmites australis and if the restoration of the native vegetation in brackish Spartina alterniflora marshes results in the reestablishment of the arthropod community. The invasion of Phragmites into a coastal Spartina marsh in southern New Jersey seriously altered arthropod assemblages and trophic structure by changing the abundance of trophic groups (detritivores, herbivores, carnivores) and their taxonomic composition. Herbivore assemblages shifted from the dominance of external free‐living specialists (e.g., planthoppers) in Spartina to concealed feeders in Phragmites (stem‐feeding cecidomyiids). Moreover, free‐living arthropods in Phragmites became dominated by detritivores such as Collembola and chironomids. The dominant marsh spiders, web‐building linyphiids, were significantly reduced in Phragmites habitats, likely caused by differences in the physical environment of the invaded habitats (e.g., lower stem densities). Thus, trophic structure of arthropod assemblages in Phragmites, as seen in the large shifts in feeding guilds, was significantly different from that in Spartina. Removal of Phragmites with the herbicide glyphosate resulted in the rapid return of Spartina (≤5 yrs). Moreover, return of the dominant vegetation was accompanied by the recovery of most original habitat characteristics (e.g., live and dead plant biomass, water flow rate). The arthropod assemblage associated with Spartina also quickly returned to its preinvasion state and was not distinguishable from that in uninvaded Spartina reference sites. This study provides evidence that the reestablishment of native vegetation in areas previously altered by an invasive plant can result in the rapid recovery of the native arthropod assemblage associated with the restored habitat.     

互花米草群落功率最大化倾向

植物新种进入新区域后,通过扰动或搅局,改变生态系统的结构,形成有特色的自组 织生态系统,趋向新条件下的功率最大化.本文应用能值分析的方法,分别对苏北互花米草 生态系统和光滩生态系统进行能值分析和系统评估.结果表明：互花米草生态系统每年能值 产出比光滩生态系统高1.52E+18 sej.能值密度是光滩的4.72倍,基础能值产出率约为光滩生态系统的5倍.互花米草生态系统能更有效地利用能量,增加系统内部的 能值贮存,实现系统内部的功率最大化.互花米草入侵某区域滩涂后,通过自组织促使整个互花米草生态系统趋于功率最大化,充分发挥生态服务功能,但由于其繁殖扩散过快,导致种群爆发,产生了一系列负面效应,如抑制本土物种、侵占航道、危害贝类养殖等.     

Effect of Salt Stress on Seed Germination and EmbryoGrowth of Spartina alterniflora (Gramineae)

In this paper , the effect of NaCl concentration ( 1× 104 - 6× 104 mg/L) on the seed germination of smooth cordgrass ( Spartina alterniflora) was investigated using 1/2 Hoagland as the basic culture solution. The results indicated that the coleoptile and embryonal axis grew before the radicle and plumule in the stage of germination . Salinity level had no influence on the seed germination percentage when the salt concentration didn′t exceed 3×104 mg/L. The optimal salinity level for seed germination and embryo growth was 1×104 mg/L . The inhibit effect of salt concentration on the growth of embryo′s different part was different, and the length of radicle and plumule decreased significantly with the increase of salt concentration. However , the lesser inhibition on the embryonal axis and coleoptile was observed under the higher salt concentration, and thus the seedlings out of soil were ensured and the species was quickly adapted to the environment of inter- tidal zone .     

Enumeration and Localization of N2-Fixing Bacteria Associated with Roots of Spartina alterniflora Loisel

Numbers and possible locations of N₂-fixing bacteria were investigated in roots of Spartina alterniflora Loisel, which support nitrogenase activity in the undisturbed native habitat. N₂-fixing bacteria were recovered in cultures both from S. alterniflora roots and from the surrounding sediment, and they formed a greater proportion of the bacteria recovered from root homogenates than from salt-marsh sediment. N₂-fixing bacteria were recovered in high numbers from the rhizoplane of S. alterniflora after roots were treated with 1 or 5% chloramine-T for 1 h or with 1% NaOCl for 1 or 2 h. Immersing S. alterniflora roots in 5% NaOCl for 1 h was more effective in distinguishing bacteria inside the roots since this treatment nearly eliminated N₂-fixing bacteria recoverable from the rhizoplane, although high numbers of N₂-fixing bacteria were recovered from homogenates of roots treated with 5% NaOCl for 1 h. However, this treatment was less effective with roots of Zea mays L. (Funks G4646) and Sorghum bicolor (L.) Moench (CK-60 A), indicating that techniques to surface sterilize roots should be evaluated for different plants. Bacteria were observed by light and electron microscopy inter- and intracellularly in the cortex and in the aerenchyma of S. alterniflora roots. This study clearly shows that bacteria, including N₂ fixers, colonize the interior of roots of S. alterniflora growing in a Chesapeake Bay, Maryland, salt marsh.