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。     

Populations of Anaerobic Phototrophic Bacteria in a Spartina alterniflora Salt Marsh

Habitat-simulating media were used with the Hungate anaerobic roll tube technique to enumerate culturable anaerobic photosynthetic bacteria in sediment, tidal waters, and Spartina alterniflora plant samples collected from the salt marsh at Sapelo Island, Ga. No phototrophs were detected in samples of creekside (low marsh) sediment or in tidal waters in creekside regions. In the high marsh region, 90% of anaerobic phototrophic bacteria occurred in the top 5 mm of sediment and none were detected below 6 mm. There was a seasonal variation, with maximal populations occurring in summer and fall (mean, 4.4 × 10⁵ phototrophs g of dry sediment⁻¹) and minimal numbers occurring in winter (mean, 3.9 × 10³ phototrophs g of dry sediment⁻¹). During winter and late spring, phototrophs had a patchy distribution over the high marsh sediment surface. In contrast, during late summer they had a random uniform distribution. Tidal water collected over high marsh sediment contained an average of 8.7 × 10² phototrophs ml⁻¹, with no significant seasonal variation. Anaerobic phototrophic bacteria were also cultured from the lower stem tissue of S. alterniflora growing in both the high (4.3 × 10⁴ phototrophs g of dry tissue⁻¹) and creekside (4.9 × 10⁴ phototrophs g of dry tissue⁻¹) marsh regions. Chromatium buderi, Chromatium vinosum, Thiospirillum sanguineum, Rhodospirillum molischianum, and Chlorobium phaeobacteroides were the predominant anaerobic phototrophic species cultured from high marsh sediment. The two Chromatium species were dominant.     

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.     

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.     

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.     

Evidence for Hygrometric Pressurization in the Internal Gas Space of Spartina alterniflora

Higher pressure, up to several hundred pascal relative to ambient, is generated by hygrometric pressurization within the central hollow space of the stem in Spartina alterniflora. Dilution of oxygen and nitrogen by water vapor within the plant's internal gas space results in an influx of nitrogen and oxygen from the air and a net increase in the internal gas pressure at steady state. The nature of the pressure gradient suggests that small pores exist in the plant tissues. Moreover, the compact arrangement of leaf mesophyll cells creates a high resistance for the mass flow of gases and contributes to the higher pressure within leaves. After experimentally venting the internal pressure, outside air diffused through the basal area of the adaxial side of the leaves into the internal space and elevated pressure was restored.     

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     

Endorhizal and Exorhizal Acetylene-reducing Activity in a Grass (Spartina alterniflora Loisel.)-Diazotroph Association

Earlier studies indicated that bacteria responsible for nitrogenase activity of some grasses are located inside the roots. Those studies were conducted with excised roots in which a long, unexplained “lag phase” occurred before initiation of nitrogenase activity. When hydroponically maintained Spartina alterniflora Loisel. was incubated in a two-compartment system with acetylene, ethylene was produced following, at most, a 2-hour lag in both the upper (shoot) and lower (roots + water) phases. Ethylene production in the upper phase not attributable to leaf-associated acetylene-reducing activity or to diffusion of ethylene from around the roots is considered to represent “endorhizal acetylene-reducing activity,” the internally produced ethylene diffusing into the upper phase via the lacunae. Ethylene produced in the lower phase is designated “exorhizal acetylene-reducing activity.” The endorhizal acetylene-reducing activity, in comparison to exorhizal activity, was relatively insensitive to additions of HgCl₂, NH₄Cl, or carbon sources to the lower phase. Post-lag acetylene-reducing activity of roots excised from plants growing in soil responded to additions in a manner similar to that of endorhizal acetylene-reducing activity, whereas post-lag acetylene-reducing activity of rhizosphere soil responded in a manner similar to that of exorhizal acetylene-reducing activity.     

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     

Climate Drivers of Spartina alterniflora Saltmarsh Production in Georgia,USA

Tidal wetlands are threatened by global changes related not only to sea level rise but also to altered weather patterns. To predict consequences of these changes on coastal communities, it is necessary to understand how temporally varying abiotic conditions drive wetland production. In 2000–2011, we conducted annual surveys of Spartina alterniflora biomass in tidal marshes at nine sites in and around the Altamaha river estuary on the coast of Georgia, USA. End of the year live biomass was assessed in the creekbank and midmarsh zones to estimate annual net primary production (ANPP). River discharge was the most important driver of S. alterniflora ANPP, especially in creekbank vegetation. Increased river discharge reduces water column salinity, and this was most likely the proximate driver of increased production. In the midmarsh zone, the patterns were less distinct, although river discharge was again the best predictor, but maximum temperature had similar predictive ability. In contrast to results from terrestrial grasslands, we found no consistent evidence for a sharply delimited critical period for any climate driver in the tidal marsh, which indicates that plant growth was responsive to abiotic drivers at any time during the growing season. Results were broadly consistent across multiple sites within a geographic region. Our results differ from previous analyses of production in S. alterniflora marshes, which either identified oceanic drivers of S. alterniflora production or were unable to identify any drivers, likely because the low-latitude sites we studied were hotter and more affected by river discharge than those in previous studies.     

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...     

Molecular Analysis of Diazotroph Diversity in the Rhizosphere of the Smooth Cordgrass, Spartina alterniflora

N₂ fixation by diazotrophic bacteria associated with the roots of the smooth cordgrass, Spartina alterniflora, is an important source of new nitrogen in many salt marsh ecosystems. However, the diversity and phylogenetic affiliations of these rhizosphere diazotrophs are unknown. Denaturing gradient gel electrophoresis (DGGE) of PCR-amplified nifH sequence segments was used in previous studies to examine the stability and dynamics of the Spartina rhizosphere diazotroph assemblages in the North Inlet salt marsh, near Georgetown, S.C. In this study, plugs were taken from gel bands from representative DGGE gels, the nifH amplimers were recovered and cloned, and their sequences were determined. A total of 59 sequences were recovered, and the amino acid sequences predicted from them were aligned with sequences from known and unknown diazotrophs in order to determine the types of organisms present in the Spartina rhizosphere. We recovered numerous sequences from diazotrophs in the γ subdivision of the division Proteobacteria (γ-Proteobacteria) and from various anaerobic diazotrophs. Diazotrophs in the α-Proteobacteria were poorly represented. None of the Spartina rhizosphere DGGE band sequences were identical to any known or previously recovered environmental nifH sequences. The Spartina rhizosphere diazotroph assemblage is very diverse and apparently consists mainly of unknown organisms.     

Molecular analysis of diazotroph diversity in the rhizosphere of the smooth cordgrass, Spartina alterniflora

N(2) fixation by diazotrophic bacteria associated with the roots of the smooth cordgrass, Spartina alterniflora, is an important source of new nitrogen in many salt marsh ecosystems. However, the diversity and phylogenetic affiliations of these rhizosphere diazotrophs are unknown. Denaturing gradient gel electrophoresis (DGGE) of PCR-amplified nifH sequence segments was used in previous studies to examine the stability and dynamics of the Spartina rhizosphere diazotroph assemblages in the North Inlet salt marsh, near Georgetown, S.C. In this study, plugs were taken from gel bands from representative DGGE gels, the nifH amplimers were recovered and cloned, and their sequences were determined. A total of 59 sequences were recovered, and the amino acid sequences predicted from them were aligned with sequences from known and unknown diazotrophs in order to determine the types of organisms present in the Spartina rhizosphere. We recovered numerous sequences from diazotrophs in the gamma subdivision of the division Proteobacteria (gamma-Proteobacteria) and from various anaerobic diazotrophs. Diazotrophs in the alpha-Proteobacteria were poorly represented. None of the Spartina rhizosphere DGGE band sequences were identical to any known or previously recovered environmental nifH sequences. The Spartina rhizosphere diazotroph assemblage is very diverse and apparently consists mainly of unknown organisms.     

Spartina alterniflora loisel root structure and meiofaunal abundance

Meiofauna were collected from adjacent areas with and without Spartina alterniflora Loisel culms. Environmental complexity resulting from root structure cannot be determined by simply observing S. alterniflora culm distribution. There were no significant differences between the two areas with respect to root density or root sizes; however, the number of live roots was significantly higher in the area containing Spartina culms. Nematodes, the most abundant taxon were positively correlated with live root density. In contrast, the number of dead roots and harpacticoid copepods were higher in the area without culms. This study suggests that nematodes are positively associated with micro-oxygenated zones produced by live S. alterniflora roots. Other measures of biogenic structure, such as total root density or root size fractions are not important in affecting nematode abundance.     

Genotypic diversity enhances invasive ability of Spartina alterniflora

Although genetic diversity is very important for alien species, which have to cope with new environments, little is known about the role that genetic diversity plays in their invasive success. In this study, we set up a manipulation experiment including three levels of genotypic diversity to test whether genotypic diversity can enhance the invasive ability of alien species, in our case the invasive Spartina alterniflora in China, and to infer the underlying mechanisms. There was no significant relationship between genotypic diversity and parameters of performance in the first year; however, from the summer of the second year onwards, genotypic diversity enhanced four of the six parameters of performance. After two growing seasons, there were significant positive relationships between genotypic diversity and maximum spread distance, patch size, shoot number per patch, and aboveground biomass. Moreover, abundance of the native dominant species Scirpus mariqueter was marginally significantly decreased with genotypic diversity of S. alterniflora, suggesting that enhanced invasive ability of S. alterniflora may have depressed the growth of the native species. There was no significant difference in most measures of performance among six genotypes, but we observed a transgressive over performance in four measures in multiple‐genotype patches. At the end of the experiment, there were significant nonadditive effects of genotypic diversity according to Monte Carlo permutations, in six‐genotype, but not three‐genotype plots. Our results indicated that both additive and nonadditive effects played roles in the positive relationship between genetic diversity and invasion success, and nonadditive effects were stronger as duration increased.     

The macrobenthos in Spartina alterniflora salt marshes of the Wanggang tidal-flat,Jiangsu coast,China

On the Jiangsu coast, eastern China, Spartina alterniflora, which was introduced artificially into the region, is becoming a dominant plant species in the inter-tidal salt marshes. In order to evaluate the environmental and ecological impact of the colonization of S. alterniflora, we carried out investigations into the benthic macrofauna of the Spartina marshes of the Wanggang area, central Jiangsu coast, in 2006 and 2007. Based on analysis of the data sets obtained, 12 species of macrobenthos have been identified for the Wanggang salt marsh, including S. alterniflora, Cerithidea cingulata, Littorna scabra, Bithynia fuchsiana, Macrophthalmus japonicus, Uca arcuata, Nereis sp., Boleophthalmus petinirostris, Cyclina sinensis, Bullacta exarata, Angustassiminea castanea and Glaucomya chinensis. The results indicate that some of the native species have adapted to the new ecological environment associated with the cordgrass S. alterniflora. The biomass of macrobenthos varies significantly over different parts of the salt marsh. Further, there is a seasonal change in bio-density, with the density in summer (July and August) > autumn (November) > early summer (May). At the landward edge and over the central part of the S. alterniflora marsh, the bio-diversity is higher than the other areas of the marsh.     

Effects of Spartina alterniflora Invasion on Soil Respiration in the Yangtze River Estuary,China

Many studies have found that plant invasion can enhance soil organic carbon (SOC) pools, by increasing net primary production (NPP) and/or decreased soil respiration. While most studies have focused on C input, little attention has been paid to plant invasion effects on soil respiration, especially in wetland ecosystems. Our study examined the effects of Spartina alterniflora invasion on soil respiration and C dynamics in the Yangtze River estuary. The estuary was originally occupied by two native plant species: Phragmites australis in the high tide zone and Scirpus mariqueter in the low tide zone. Mean soil respiration rates were 185.8 and 142.3 mg CO₂ m⁻² h⁻¹ in S. alterniflora and P. australis stands in the high tide zone, and 159.7 and 112.0 mg CO₂ m⁻² h⁻¹ in S. alterniflora and S. mariqueter stands in the low tide zone, respectively. Aboveground NPP (ANPP), SOC, and microbial biomass were also significantly higher in the S. alterniflora stands than in the two native plant stands. S. alterniflora invasion did not significantly change soil inorganic carbon or pH. Our results indicated that enhanced ANPP by S. alterniflora exceeded invasion-induced C loss through soil respiration. This suggests that S. alterniflora invasion into the Yangtze River estuary could strengthen the net C sink of wetlands in the context of global climate change.     

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.     

Relative Importance of Ion Exclusion, Secretion and Accumulation in Spartina alterniflora Loisel.

The extent to which Spartina alterniflora Loisel. excluded,secreted or accumulated the major seawater ions (Cl^-, SO^2-₄,Na⁺, K⁺, Mg²⁺, and Ca²⁺) was investigated under varying salinitytreatments. From a quantitative viewpoint, ion exclusion wasmost prominent and accounted for 91–97% of the theoreticalmaximum ion uptake as a result of transpiration and growth.Of those ions taken up, approximately half was secreted fromthe shoots. Relative to K⁺, a disproportionate amount of Na⁺was excluded at the roots and secreted by the shoots. The concentrationwithin the tissues of S. alterniflora did not change with salinitytreatment for the majority of the ions examined, but Na⁺ wasmore than twice as concentrated at 40 g dm^-3 than at lOgdm^-3.Calculations of the flux of ions from salt marsh sediments tothe flood water via shoot secretion or stem/leaf turnover indicatethat these processes may be important to the ecology of S. alternifloraas mechanisms that limit the accumulation of salt within theroot zone.     

Interaction among sediment anaerobiosis, nitrogen uptake and photosynthesis of Spartina alterniflora

Spartina alterniflora Lois. plants were grown under controlled sediment-pH-redox conditions. Uptake of added ¹⁵NH₄-nitrogen and subsequent photosynthetic activity under different redox conditions in the plant root rhizosphere were measured. Data for total plant nitrogen and ¹⁵N concentration indicated that nitrogen allocation was not altered by anaerobic conditions of the sediment. However, average net photosynthesis was reduced by up to 35% for plants under anaerobiosis. The results indicate that anaerobiosis in the root rhizosphere, rather than limiting nitrogen uptake, influences photosynthesis and growth of S. alterniflora under anaerobic conditions.