Small-scale resource heterogeneity among halophytic plant species in an upper salt marsh community

The focus of the present study was to determine whether spatial and temporal variation, in soil and plant factors of upper salt marsh areas, exists among microsites of three coexisting halophytic plant species, Grindelia humilis, Jaumea carnosa and Limonium californicum, on the Pacific coasts of California. Low frequencies in tidal inundations coupled with variation in temperature and rainfall patterns often result in wide seasonal salinity variations in these upper salt marsh zones. The study was carried out to evaluate (1) variation in soil osmolalities and soil water contents for the three species; (2) differences in chemical soil factors associated with the three species; (3) variation in leaf succulence associated with saline stress; and (4) levels of plant nutrients present in each of the three species. The results indicated that soil osmolalities associated with Limonium, a salt excreting halophyte were markedly different from those recorded for the other two species. Nitrogen levels in soils associated with Jaumea as well as levels in composite plant samples were significantly lower than those of the other two species. Also noted were variations in nutrient ions in plants as well as soil environments.     

Crenarchaeal heterotrophy in salt marsh sediments

Mesophilic Crenarchaeota (also known as Thaumarchaeota) are ubiquitous and abundant in marine habitats. However, very little is known about their metabolic function in situ. In this study, salt marsh sediments from New Jersey were screened via stable isotope probing (SIP) for heterotrophy by amending with a single ¹³C-labeled compound (acetate, glycine or urea) or a complex ¹³C-biopolymer (lipids, proteins or growth medium (ISOGRO)). SIP incubations were done at two substrate concentrations (30–150 μM; 2–10 mg ml⁻¹), and ¹³C-labeled DNA was analyzed by terminal restriction fragment length polymorphism (TRFLP) analysis of 16S rRNA genes. To test for autotrophy, an amendment with ¹³C-bicarbonate was also performed. Our SIP analyses indicate salt marsh crenarchaea are heterotrophic, double within 2–3 days and often compete with heterotrophic bacteria for the same organic substrates. A clone library of ¹³C-amplicons was screened to find matches to the ¹³C-TRFLP peaks, with seven members of the Miscellaneous Crenarchaeal Group and seven members from the Marine Group 1.a Crenarchaeota being discerned. Some of these crenarchaea displayed a preference for particular carbon sources, whereas others incorporated nearly every ¹³C-substrate provided. The data suggest salt marshes may be an excellent model system for studying crenarchaeal metabolic capabilities and can provide information on the competition between crenarchaea and other microbial groups to improve our understanding of microbial ecology.     

Dimethyl sulfide metabolism in salt marsh sediments

Abstract Anoxic sediment slurries prepared from Spartina salt marsh soils contained dimethyl sulfide (DMS) at concentrations ranging from 1 to 10 μM. DMS was produced in slurries over the initial 1–24 h incubation. After the initial period of production, DMS decreased to undetectable levels and methane thiol (MSH) was produced. Inhibition of methanogenesis caused a 20% decrease in the rate of DMS consumption, while inhibition of sulfate reduction caused a 80% decrease in DMS consumption. When sulfate reduction and methanogenesis were simultaneously inhibited, DMS did not decrease. DMS contributed about 28% to the methane production rate, while DMS probably contributed only 1% or less to the sulfate reduction rate. Incubation of the sediment slurries under an atmosphere of air resulted in similar DMS consumption compared to anaerobic incubations, but MSH and CH₄ were not evolved.
Sediments from the marsh released significant quantities of DMS when treated with cold alkali, indicating that potentially significant sources of DMS existed in the sediments. Values of base-hydrolyzable DMS as high as 190 μmol per liter of sediment were observed near the sediment surface, and values always decreased with depth in the sediment. Simple flux experiments with small intact sediment cores, showed that DMS was emitted from the marsh surface when cores were injected with glutaraldehyde or molybdate and 2-bromoethanesulfonate (BES), but nit when cores were left uninhibited. These results showed that DMS was readily metabolized by microbes in marsh sediments and that this metabolism may be responsible for reducing the emission of DMS from the marsh surface.     

Nitrogen resorption from senescing leaves of three salt marsh plant species

Seasonalvariation in leaf nitrogen of mature green and senescent leaves and nitrogenresorption efficiency in three plants (Spartina maritima, Halimioneportulacoides and Arthrocnemum perenne) of aTagus estuary salt marsh are reported. Total nitrogen concentrations in greenand senescent leaves were higher during winter (December and March). Soilinorganic nitrogen availability showed an opposite pattern with higherconcentrations during summer (June and September) when total leaf biomass washigher. Nitrogen resorption efficiency ranged between 31 and 76% andH. portulacoides was the plant that better minimizednitrogen loss by this process. Nitrogen resorption occurred mainly from thesoluble protein pool, although other fractions must have been broken down duringthe resorption process. No significant seasonal variation in nitrogen resorptionefficiency and no relation to leaf total nitrogen or soil nitrogen availabilitywere found. This suggests that the efficiency of the resorption process is notdetermined by the plant nitrogen status nor by the availability of the nutrientin the soil. Nevertheless, resorption from senescing leaves may play animportant role in the nitrogen dynamics of salt marsh plants and reduce thenitrogen requirements for plant growth.     

Microbial carbon monoxide consumption in salt marsh sediments

We have examined sediments from a fringing salt marsh in Maine to further understand marine CO metabolism, about which relatively little is known. Intact cores from the marsh emitted CO during dark oxic incubations, but emission rates were significantly higher during anoxic incubations, which provided evidence for simultaneous production and aerobic consumption in surface sediments. CO emission rates were also elevated when cores were exposed to light, which indicated that photochemical reactions play a role in CO production. A kinetic analysis of marsh surface sediments yielded an apparent K(m) of about 82 ppm, which exceeded values reported for well-aerated soils that consume atmospheric CO (65nM). Surface (0-0.2 cm depth interval) sediment slurries incubated under oxic conditions rapidly consumed CO, and methyl fluoride did not inhibit uptake, which indicated that neither ammonia nor methane oxidizers contributed to the observed activity. In contrast, aerobic CO uptake was inhibited by additions of readily available organic substrates (pyruvate, glucose and glycine), but not by cellulose. CO was also consumed by surface and sub-surface sediment slurries incubated under anaerobic conditions, but rates were less than during aerobic incubations. Molybdate and nitrate or nitrite, but not 2-bromoethanesulfonic acid, partially inhibited anaerobic uptake. These results suggest that sulfidogens and acetogens, but not dissimilatory nitrate reducers or methanogens, actively consume CO. Sediment-free plant roots also oxidized CO aerobically; rates for Spartina patens and Limonium carolinianum roots were significantly higher than rates for Spartina alterniflora roots. Thus plants may also impact CO cycling in estuarine environments.     

Microbial indicators of oil-rich salt marsh sediments

Selected microbial parameters were monitored in sediments from a pristine and an oil-field salt marsh. Although numbers of hydrocarbonoclastic bacteria and fungi were significantly greater in the oil field, the values did not show a strong correlation with levels of hydrocarbons (r = 0.43 and r = 0.49, respectively). However, a high correlation was noted between ratios of hydrocarbonoclastic and total aerobic heterotrophic bacteria and levels of hydrocarbons as well as the relative concentration of hydrocarbons (ratio of hydrocarbons to chloroform extractables) (r = 0.87 and r = 0.77, respectively). Data suggest that this first ratio is a more valid microbial indicator of hydrocarbon abundance than other factors examined. Significant differences in the ratio of pigmented to total colony-forming units, the ratio of different to total colony-forming units, and the diversity index were noted between the natural and oil-field marsh. It is suggested that the presence of hydrocarbons alters the relative abundance of the most predominant aerobic heterotrophic bacteria.     

Interactions between C3 and C4 salt marsh plant species during four years of exposure to elevated atmospheric CO2

Elevated atmospheric CO2 is known to stimulate photosynthesis and growth of plants with the C₃ pathway but less of plants with the C₄ pathway. An increase in the CO₂ concentration can therefore be expected to change the competitive interactions between C₃ and C₄ species. The effect of long term exposure to elevated CO₂ (ambient CO₂ concentration +340 µmol CO₂ mol^-1) on a salt marsh vegetation with both C₃ and C₄ species was investigated. Elevated CO₂ increased the biomass of the C₃ sedgeScirpus olneyi growing in a pure stand, while the biomass of the C₄ grassSpartina patens in a monospecific community was not affected. In the mixed C₃/C₄ community the C₃ sedge showed a very large relative increase in biomass in elevated CO₂ while the biomass of the C₄ species declined.The C₄ grassSpartina patens dominated the higher areas of the salt marsh, while the C₃ sedgeScirpus olneyi was most abundant at the lower elevations, and the mixed community occupied intermediate elevations.Scirpus growth may have been restricted by drought and salt stress at the higher elevations, whileSpartina growth at the lower elevations may be affected by the higher frequency of flooding. Elevated CO₂ may affect the species distribution in the salt marsh if it allowsScirpus to grow at higher elevations where it in turn may affect the growth ofSpartina.     

Methanogenesis and microbial lipid synthesis in anoxic salt marsh sediments

In anoxic salt marsh sediments of Sapelo Island, GA, USA, the vertical distribution of CH₄ production was measured in the upper 20 cm of surface sediments in ten locations. In one section of high marsh sediments, the concentration and oxidation of acetate in sediment porewaters and the rate and amount of¹⁴C acetate and¹⁴CO₂ incorporation into cellular lipids of the microbial population were investigated. CH₄ production rates ranged from <1 to 493 nM CH₄ gram sediment⁻¹ day⁻¹ from intact subcores incubated under nitrogen. Replacement with H₂ stimulated the rate of methane release up to nine fold relative to N₂ incubations. Rates of lipid synthesis from CO₂ averaged 39.2 ×10⁻²nanomoles lipid carbon cm³ sediment⁻¹ hr⁻¹, suggesting that CO₂ may be an important carbon precursor for microbial membrane synthesis in marsh sediments under anoxic conditions. Qualitative measurements of lipid synthesis rates from acetate were found to average 8.7 × 10⁻² nanomoles. Phospholipids were the dominant lipids synthesized by both substrates in sediment cores, accounting for an average of 76.6% of all lipid radioactivity. Small amounts of ether lipids indicative of methanogenic bacteria were observed in cores incubated for 7 days, with similar rates of synthesis for both CO₂ and acetate. The low rate of ether lipid synthesis suggests that either methanogen lipid biosynthesis is very slow or that methanogens represent a small component of total microbial lipid synthesis in anoxic sediments. present address: The University of Maryland,, Chesapeake Biological Laboratory, Box 38, Solomons, MD 20688, USA     

Methanogenesis and microbial lipid synthesis in anoxic salt marsh sediments

In anoxic salt marsh sediments of Sapelo Island, GA, USA, the vertical distribution of CH₄ production was measured in the upper 20 cm of surface sediments in ten locations. In one section of high marsh sediments, the concentration and oxidation of acetate in sediment porewaters and the rate and amount of¹⁴C acetate and¹⁴CO₂ incorporation into cellular lipids of the microbial population were investigated. CH₄ production rates ranged from <1 to 493 nM CH₄ gram sediment⁻¹ day⁻¹ from intact subcores incubated under nitrogen. Replacement with H₂ stimulated the rate of methane release up to nine fold relative to N₂ incubations. Rates of lipid synthesis from CO₂ averaged 39.2 ×10⁻²nanomoles lipid carbon cm³ sediment⁻¹ hr⁻¹, suggesting that CO₂ may be an important carbon precursor for microbial membrane synthesis in marsh sediments under anoxic conditions. Qualitative measurements of lipid synthesis rates from acetate were found to average 8.7 × 10⁻² nanomoles. Phospholipids were the dominant lipids synthesized by both substrates in sediment cores, accounting for an average of 76.6% of all lipid radioactivity. Small amounts of ether lipids indicative of methanogenic bacteria were observed in cores incubated for 7 days, with similar rates of synthesis for both CO₂ and acetate. The low rate of ether lipid synthesis suggests that either methanogen lipid biosynthesis is very slow or that methanogens represent a small component of total microbial lipid synthesis in anoxic sediments. present address: The University of Maryland,, Chesapeake Biological Laboratory, Box 38, Solomons, MD 20688, USA     

Microbial activity profiles in Tagus estuary salt marsh sediments

This study provides some results about microbial activity in salt marsh sediments. Microbial activity was determined by profiling extracellular enzyme activities in three Tagus estuary marshes and in two sediments horizons: surface layer (0–2 cm) and depth (8–10 cm). Five enzymatic activities were examined (β-glucosidase, cellulase, alkaline phosphatase, potential nitrification and nitrate reductase). All extracellular enzymatic activities were highest in the surface layer and decreased with depth. β-glucosidase and alkaline phosphatase prevailed both in surface sediments (1150 and 1200 ηmol h⁻¹ g⁻¹, respectively) and in deeper sediments (150 and 200 ηmol h⁻¹ g⁻¹, respectively). Microbial activities differed significantly between salt marshes. The marsh location in the estuary seemed to contribute to these differences: marshes located in the proximity of urbanised and industrial areas had higher microbial activities.     

Denitrification in salt marsh sediments: Evidence for seasonal temperature selection among populations of denitrifiers

Direct measurements of bacterial denitrification in salt marsh sediments near Woods Hole, Massachusetts were made over a 10-month period using a simple and precise gas-chromatographic technique. Based on laboratory experiments at 5°, 10°, and 20°C, it is shown that seasonal temperature variations select for at least two distinct populations of denitrifiers.In situ incubations suggest that resident populations of denitrifying bacteria are cold-sensitive. Salt marsh denitrifying bacteria are not optimally adapted to their thermal environment, but to temperatures 5°–10°C higher. In these water-logged muds, rates of bacterial denitrification (0.3–1.5g N₂/gm sediment-hr) are up to three orders of magnitude greater than maximum potential rates of insitu bacterial and algal nitrogen fixation.Supported by the Victoria Foundation and NSF grants GA 28365, GA 28272, and GA 41506. This is Contribution No. 3730 from the Woods Hole Oceanographic Institution.     

Interactions among salt marsh plants vary geographically but not latitudinally along the California coast

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.     

The microbially driven formation of siderite in salt marsh sediments

We employ complementary field and laboratory‐based incubation techniques to explore the geochemical environment where siderite concretions are actively forming and growing, including solid‐phase analysis of the sediment, concretion, and associated pore fluid chemistry. These recently formed siderite concretions allow us to explore the geochemical processes that lead to the formation of this less common carbonate mineral. We conclude that there are two phases of siderite concretion growth within the sediment, as there are distinct changes in the carbon isotopic composition and mineralogy across the concretions. Incubated sediment samples allow us to explore the stability of siderite over a range of geochemical conditions. Our incubation results suggest that the formation of siderite can be very rapid (about two weeks or within 400 hr) when there is a substantial source of iron, either from microbial iron reduction or from steel material; however, a source of dissolved iron is not enough to induce siderite precipitation. We suggest that sufficient alkalinity is the limiting factor for siderite precipitation during microbial iron reduction while the lack of dissolved iron is the limiting factor for siderite formation if microbial sulfate reduction is the dominant microbial metabolism. We show that siderite can form via heated transformation (at temperature 100°C for 48 hr) of calcite and monohydrocalcite seeds in the presence of dissolved iron. Our transformation experiments suggest that the formation of siderite is promoted when carbonate seeds are present.     

Spatial and temporal variations in salt marsh microorganisms of the Wadden Sea

Salt marshes exist at the interface of the marine and the terrestrial system. Shore height differences and associated variations in inundation frequency result in altered abiotic conditions, plant communities, and resource input into the belowground system. These factors result in three unique zones, the upper salt marsh (USM), the lower salt marsh (LSM), and the pioneer zone (PZ). Marine detritus, such as micro‐ and macroalgae, is typically flushed into the PZ daily, with storm surges moving both salt marsh detritus and marine detritus into higher salt marsh zones. Microbial assemblages are essential for the decomposition of organic matter and have been shown to sensitively respond to changes in abiotic conditions such as oxygen supply and salinity. However, temporal and spatial dynamics of microbial communities of Wadden Sea salt marshes received little attention. We investigated the dynamics of soil microbial communities across horizontal (USM, LSM, and PZ), vertical (0–5 and 5–10‐cm sediment depth), and temporal (spring, summer, and autumn) scales in the Wadden Sea salt marsh of the European North Atlantic coast using phospholipid fatty acid (PLFA) analysis. Our results show strong spatial dynamics both among salt marsh zones and between sediment depths, but temporal dynamics to be only minor. Despite varying in space and time, PLFA markers indicated that bacteria generally were the dominant microbial group across salt marsh zones and seasons, however, their dominance was most pronounced in the USM, whereas fungal biomass peaked in the LSM and algal biomass in the PZ. Only algal markers and the stress marker monounsaturated to saturated fatty acid ratio responded to seasonality. Overall, therefore, the results indicate remarkable temporal stability of salt marsh microbial communities despite strong variability in abiotic factors.     

The effect of competition and salinity on the growth of a salt marsh plant species

Summary Young rhizome sprouts of the herbaceous perennial Jaumea carnosa were propagated from material collected in a salt marsh along the central California coast. The sprouts were transplanted to flats of sand sown with different densities of seeds of a representative glycophyte, Lolium perenne L. Derby, turf type. Controls flats contained only Jaumea or Lolium. Three series of replicated flats were watered from above with dilutions of seawater in 1/10 strength Hoagland solution, such that dissolved salts were 400, 4000 or 11,600 ppm. Two other series were continuously subirrigated with 400 or 11,600 ppm salt water. After 61 days of treatment in a greenhouse with a 30/11°C thermoperiod (mean daily max/min), all plants were harvested and weighed. In the monospecific control flats, the growth of both species declined with increasing salinity, but the relative decline of Lolium was three times that of Jaumea. Jaumea's root: shoot ratio was also less affected by salinity. Both species grew well when subirrigated by 400 ppm salt water, but grew poorly when subirrigated by 11,600 ppm salt water, indicating that aeration alone is not the most significant factor in the marsh. The effect of interspecific competition on Jaumea was marked at low salinity, depressing growth by 52% compared to controls, but at high salinity the competitive effect was insignificant, whether the plants were watered from above or subirrigated. This supports the hypothesis that intolerant halophytes such as Jaumea are restricted in nature to salt marshes because they are poor competitors with glycophytes on non-saline soils.This research was supported by the John Simon Guggenheim Memorial Foundation and by the University of California Bodega Marine Laboratory     

Interactions of plant species mediated plant competition for inorganic nitrogen with soil microorganisms in an alpine meadow

The aim of the present work was to evaluate the effects of regulated deficit irrigation (RDI) applied in the post-harvest stage of peach trees. The 3-year trial was carried out in Italy (N 40°20′, E 16°48′) on mature peach plants (cv “Springcrest”) trained to transverse Y. From bud break to harvest, irrigation was carried out by applying 100% ET_c, while from harvest to early autumn, plants were separated into three groups and subjected to different irrigation treatments (100, 57 and 34% ET_c). The decrease in soil water content caused a reduction in the values of tissue water potential and gas exchange both in 57% ET_c and 34% ET_c treatments. RDI determined the reduction in the growth of waterspouts and lateral shoots but did not influence the growth of fruiting shoots. During the trial, no significant reductions in crop yield and quality were observed in the 57% ET_c treatment, whereas about 1,100, 1,800 and 2,500 m³ ha⁻¹ of water were saved in the first, the second and the third year, respectively. In the second year of the trial, the use of RDI in the post-harvest stage determined carbohydrate and nitrogen accumulation in roots, branches, shoots and floral buds. The results demonstrate that, under scarce water supply conditions, a clear benefit can be obtained through the use of RDI during the post-harvest stage. This confirms the possibility to reduce the irrigation water by applying RDI during phenological stages less sensitive to water deficit without negatively affecting peach growth and yield.     

Quantification of ammonia-oxidizing bacteria and factors controlling nitrification in salt marsh sediments

To elucidate the geomicrobiological factors controlling nitrification in salt marsh sediments, a comprehensive approach involving sediment geochemistry, process rate measurements, and quantification of the genetic potential for nitrification was applied to three contrasting salt marsh habitats: areas colonized by the tall (TS) or short (SS) form of Spartina alterniflora and unvegetated creek banks (CBs). Nitrification and denitrification potential rates were strongly correlated with one another and with macrofaunal burrow abundance, indicating that coupled nitrification-denitrification was enhanced by macrofaunal burrowing activity. Ammonia monooxygenase (amoA) gene copy numbers were used to estimate the ammonia-oxidizing bacterial population size (5.6 x 10(4) to 1.3 x 10(6) g of wet sediment(-1)), which correlated with nitrification potentials and was 1 order of magnitude higher for TS and CB than for SS. TS and CB sediments also had higher Fe(III) content, higher Fe(III)-to-total reduced sulfur ratios, higher Fe(III) reduction rates, and lower dissolved sulfides than SS sediments. Iron(III) content and reduction rates were positively correlated with nitrification and denitrification potential and amoA gene copy number. Laboratory slurry incubations supported field data, confirming that increased amounts of Fe(III) relieved sulfide inhibition of nitrification. We propose that macrofaunal burrowing and high concentrations of Fe(III) stimulate nitrifying bacterial populations, and thus may increase nitrogen removal through coupled nitrification-denitrification in salt marsh sediments.