Effects of growing conditions on the growth of and interactions between salt marsh plants: implications for invasibility of habitats

A common but often less tested explanation for the successful invasion of alien species is that invasive alien species outcompete their co-occurring natives, which may not always be the case. In this study, we established artificial environmental gradients in a series of pot experiments with controlled environments to investigate the effects of salinity, sediment type and waterlogging on the performance of and interactions between Phragmites australis (native) and Spartina alterniflora (alien), which generally co-exist in the saline intertidal zones of Chinese and American coasts. Significant effects of salinity and waterlogging were detected on biomass production and morphological characteristics of S. alterniflora and P. australis, and the competitive interactions between the two species were found to vary with all three environmental factors in our experiments. Relative Neighbor Effect (RNE) analyses indicate that competitive dominance of S. alterniflora occurred under the conditions of high salinity, sandy sediment and full immersion, whereas P. australis showed competitive dominance under the conditions of low salinity and non-immersion. Our results suggest that S. alterniflora might outcompete P. australis under conditions present in early salt marsh succession, which support the viewpoint that the outcomes of competition between co-occurring native and invasive alien plants depend on the growing conditions. The implication of this study is that in response to the environmental changes expected from seawater intrusion and sea-level rise, the range of S. alterniflora is expected to expand further in the Yangtze River estuary in the future.     

Determinants of annual–perennial plant zonation across a salt–fresh marsh interface: a multistage assessment

Vegetation zonation patterns in coastal marshes are hypothesized to be the result of both physical stress and competitive interactions. How these patterns may be driven by these factors at different life history stages remains poorly understood. We investigated the relative importance of species tolerance (response to physical stress) and competitive ability in determining the distributions of two dominant marsh species across a salt–fresh marsh interface in the Yellow River Estuary, China. There is a steep gradient in salinity across this interface and Suaeda salsa, an annual, dominates the saline side of the interface, while Phragmites australis, a perennial species, dominates the freshwater side. Using a series of field transplants, we examined the roles of physical stress and competition in mediating this zonation at different life history stages. Suaeda salsa performed well in its home zone, but seedling emergence, seedling survival, adult survival and adult growth were significantly suppressed by competition in the freshwater P. australis zone. Emergence, survival and growth of P. australis were inhibited in the saline S. salsa zone, regardless of neighbor treatments, but it performed well in its home zone. The magnitude of the competitive effect on the performance of S. salsa differed among the life history stages. Competition from P. australis had a much stronger effect on S. salsa seedling emergence and adult growth than on seedling survival and adult survival. Our results reveal that both physical stress and competition contributed to the observed zonation patterns in this marsh system. However, for S. salsa, the effect of competition varied with life-history stage. Insight into these ecological processes is critical to understanding how the zonation pattern in the marsh system is formed and maintained.     

盐度及种间相互作用对海三棱藨草、互花米草萌发及生长的影响

盐度和种间作用是影响湿地植物群落构建的关键因子。然而,已有研究主要集中于植物成体阶段,我们对生活史早期更新阶段的种间相互作用了解十分有限。崇明东滩国家级自然保护区是位于长江口的重要湿地,外来入侵植物互花米草对优势土著物种海三棱藨草的竞争排斥对当地生态系统造成了严重的负面影响。通过受控实验探讨了盐度及种间作用对海三棱藨草和互花米草种子萌发及生长的影响,以深入了解更新过程在盐沼湿地植物群落构建中的作用。结果表明,在培养皿中盐度对海三棱藨草的萌发有显著抑制作用,互花米草的萌发率受盐度影响不显著但萌发进程被延迟。混种处理对两者的萌发存在一定促进效应,且其作用强度受到盐度的调控。海三棱藨草与互花米草种子在萌发阶段的相互促进并非是通过化感作用实现的,可能是由于萌发过程对盐分的吸收减弱了盐胁迫的影响。盆栽条件下,两物种混种时的萌发及生长表现（高度、地上生物量）较单种时有所下降,但差异不显著。种间竞争受环境胁迫程度及生活史阶段的影响,竞争作用在胁迫较弱的淡水环境及生活史后期更强。盐沼湿地植物群落在形成早期受到盐度等环境因子的影响较大,不同植物对盐胁迫的响应是影响种群建成的主导因素,后期种间竞争的重要性不断增加,最终决定了植物群落的整体格局。     

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.     

Influence of Mycorrhizal Inoculation, Inundation Period, Salinity, and Phosphorus Availability on the Growth of Two Salt Marsh Grasses, Spartina alterniflora Lois. and Spartina cynosuroides (L.) Roth., in Nursery Systems

Restoration of salt marsh ecosystems is an important concern in the eastern United States to mitigate damage caused by industrial development. Little attention has been directed to the mycorrhizal influence on plantings of salt marsh species to stabilize estuarine sediments and establish cover. In our study, seedlings of two salt marsh grasses, Spartina alterniflora and Spartina cynosuroides, were grown in soil with a commercial, mixed species inoculum of arbuscular mycorrhizal fungi. Plants were grown in experimental “ebb and flow” boxes, simulating three levels of tidal inundation, to which two levels of applied phosphorus (P) and two levels of salinity were imposed. After 2.5 months, S. alterniflora was poorly colonized by arbuscular mycorrhizae, developing only fungal hyphae and no arbuscules, but S. cynosuroides became moderately colonized. Mycorrhizal inoculation marginally improved growth and P and nitrogen (N) content of both plant species at low levels of P supply but significantly increased tillering in both plant species. This factor could be beneficial in enhancing ground cover during restoration procedures. Greater P availability increased the mycorrhizal status of S. cynosuroides and improved P nutrition of both plant species, despite a reduction in the root‐to‐shoot ratio. Increasing salinity reduced mycorrhizal colonization of S. alterniflora but not of S. cynosuroides. Growth and nutrient content of S. alterniflora was improved at higher levels of salinity, but only increased nutrient content in S. cynosuroides. Increased duration of tidal inundation decreased plant growth in both species, but tissue P and N concentrations were highest with the longest time of inundation in both species.     

Edaphic diatom communities associated with Spartina alterniflora and S. patens in New Jersey

Edaphic diatoms inhabiting the sediments beneath dwarf Spartina alterniflora Loisel. and S. patens (Ait.) Muhl. in Great Bay salt marsh, Tuckerton, New Jersey were collected from 24 September 1974 through 20 August 1975. Of the 91 taxa encountered, 8 were endemic to the dwarf S. alterniflora habitat and 42 endemic to the S. patens habitat. The edaphic diatom community associated with S. patens was comprised of a much greater number of taxa and possessed higher values for species diversity (H') and evenness (J') than the community associated with dwarf S. alterniflora. The salinity of the marsh surface showed a completely opposite trend, being greatly reduced at the S. patens habitat. A highly significant relationship (P < 0.001) between the number of diatom taxa and marsh surface salinity at the S. patens habitat was demonstrated by a least squares regression. This finding led to the conclusion that the dissimilarity in the structure of the two edaphic diatom communities was primarily due to the very low marsh surface salinities at the S. patens habitat from January through June, and that this sustained. low-salinity regime allowed a very large number of taxa to coexist only in the S. patens community. Comparison of the diatom flora of Great Bay salt marsh with that of a Delaware marsh studied previously by the author showed that 67.0% of the 91 taxa encountered in New Jersey also occur on the Delaware marsh.     

Exotic Spartina alterniflora invasion alters ecosystem–atmosphere exchange of CH4 and N2O and carbon sequestration in a coastal salt marsh in China

Coastal salt marshes are sensitive to global climate change and may play an important role in mitigating global warming. To evaluate the impacts of Spartina alterniflora invasion on global warming potential (GWP) in Chinese coastal areas, we measured CH₄ and N₂O fluxes and soil organic carbon sequestration rates along a transect of coastal wetlands in Jiangsu province, China, including open water; bare tidal flat; and invasive S. alterniflora, native Suaeda salsa, and Phragmites australis marshes. Annual CH₄ emissions were estimated as 2.81, 4.16, 4.88, 10.79, and 16.98 kg CH₄ ha^?1 for open water, bare tidal flat, and P. australis, S. salsa, and S. alterniflora marshes, respectively, indicating that S. alterniflora invasion increased CH₄ emissions by 57–505%. In contrast, negative N₂O fluxes were found to be significantly and negatively correlated (P < 0.001) with net ecosystem CO₂ exchange during the growing season in S. alterniflora and P. australis marshes. Annual N₂O emissions were 0.24, 0.38, and 0.56 kg N₂O ha^?1 in open water, bare tidal flat and S. salsa marsh, respectively, compared with ‐0.51 kg N₂O ha^?1 for S. alterniflora marsh and ?0.25 kg N₂O ha^?1 for P. australis marsh. The carbon sequestration rate of S. alterniflora marsh amounted to 3.16 Mg C ha^?1 yr^?1 in the top 100 cm soil profile, a value that was 2.63‐ to 8.78‐fold higher than in native plant marshes. The estimated GWP was 1.78, ?0.60, ?4.09, and ?1.14 Mg CO₂eq ha^?1 yr^?1 in open water, bare tidal flat, P. australis marsh and S. salsa marsh, respectively, but dropped to ?11.30 Mg CO₂eq ha^?1 yr^?1 in S. alterniflora marsh. Our results indicate that although S. alterniflora invasion stimulates CH₄ emissions, it can efficiently mitigate increases in atmospheric CO₂ and N₂O along the coast of China.     

Growth strategies of four salt marsh plants on Mankyung River estuary in Korea

The emergence of seedlings, the length of roots and shoots, and the biomass of four dominant plant species and shore height were measured to investigate the growth strategy of these plants on the salt marsh of Mankyung River estuary. Four salt marsh plants showed a distinctive zonation, for example, Suaeda japonica was predominantly spread around the low salt marsh, Atriplex gmelini and Aster tripolium were in the middle, and S. asparagoides was in the upper part of the marsh. In terms of emergence of seedlings, S. japonica appeared first followed by A. gmelini, S. asparagoides, and A. tripolium. The growth strategies of halophytes were as follows: S. japonica germinated earlier than the other halophytes so that its root grew rapidly and extensively at the beginning of growth. This species adopted a continuous germination strategy, allowing growth whenever favorable conditions were provided. A. gmelini germinated later than S. japonica, as a quasi-simultaneous germination type, it showed the highest germination rate within the shortest time. Aster tripolium germinated later than any other halophyte. Since this species exhibited characteristics between the continuous germination type and the quasi-simultaneous germination type, it did not show a very high germination rate. Instead, it showed continuous germination and consistent growth of both above-ground and underground parts. Suaeda asparagoides showed an especially high emergence rate at the beginning of its growth. However, the high density retarded its growth until the middle stage. Its roots extended longer than the other halophytes, allowing it to grow well in the dry conditions of the upper marsh.     

Effect of mercury and cadmium on germination of Spartina alterniflora Loisel seeds at various salinities

Spartina alterniflora seeds were germinated in solutions containing mercury or cadmium at various salinity levels for 30 days under controlled conditions. Daily counts of epicotyl emergence were used to calculate an intial rate of germination (first 5 days), time for seeds treated with metal and salt to yield 50% of untreated seeds (distilled water treatment) and total germination at the end of the experiment (30 days). Seeds remaining ungerminated after 30 days were put through a series of washings and treatments to remove metal and salinity stresses. This was done in such a way so that osmotic and toxic effects could be identified. Results indicated both mercury and cadmium reduce the germination rate, total performance and long-term viability after an initial stimulation of germination. Increases in the detrimental effect were directly correlated with increases in salinity level and metal concentration. Findings also indicate some salt (possibly chloride ion) toxicity to S. alterniflora seeds.     

The influence of salinity on the kinetics of NH inf4 sup+ uptake in Spartina alterniflora

Summary The effects of short- and long-term exposure to a range in concentration of sea salts on the kinetics of NH _inf4 ^sup+ uptake by Spartina alterniflora were examined in a laboratory culture experiment. Long-term exposure to increasing salinity up to 50 g/L resulted in a progressive increase in the apparent K_m but did not significantly affect V_max (mean V_max=4.23±1.97 mole·g^–1·h^–1). The apparent K_m increased in a nonlinear fashion from a mean of 2.66±1.10 mole/L at a salinity of 5 g/L to a mean of 17.56±4.10 mole/L at a salinity of 50 g/L. These results suggest that the long-term effect of exposure to total salt concentrations within the range 5–50 g/L was a competitive inhibition of NH _inf4 ^sup+ uptake in S. alterniflora. No significant NH _inf4 ^sup+ uptake was observed in S. alterniflora exposed to 65 g/L sea salts. Short-term exposure to rapid changes in salinity significantly affected both V_max and K_m. Reduction of solution salinity from 35 to 5 g/L did not change V_max but reduced K_m by 71%. However, exposing plants grown at 5 g/L salinity to 35 resulted in an decrease in V_max of approximately 50%. Exposure of plants grown at 35 g/L to a total sea salt concentration of 50 g/L for 48h completely inhibited uptake of NH _inf4 ^sup+ . For both experiments, increasing salinity led to an increase in the apparent K_m similar to that found in response to long-term exposure. Our data are consistent with a conceptual model of changes in the productivity of S. alterniflora in the salt marsh as a function of environmental modification of NH _inf4 ^sup+ uptake kinetics.     

What confines an annual plant to two separate zones along coastal topographic gradients?

We investigated the roles of flooding, salinity, and plant competition in creating a bimodal zonation pattern of the marsh dominant annual plant, Suaeda salsa, along coastal topographic gradients on the Pacific coast of northern China. In two consecutive years, we manipulated salinity and flooding, salinity, and competition for S. salsa seedlings that had been transplanted into the mudflat, the high marsh, and the upland, respectively. S. salsa plants that had been transplanted into the mudflat were completely eliminated in the non-elevated treatments whereas they performed much better in the 10 cm elevated treatments, regardless of salinity treatments. Although the performance of S. salsa transplanted into the high marsh did not differ between the fresh (watered) and the salt (control) treatments, S. salsa seedling emergence in the high marsh was nearly completely inhibited in the salt treatments. In contrast, a large number of S. salsa seedlings did emerge in the fresh treatments. S. salsa transplanted into the upland performed well when neighbors were removed, whereas it appeared to be strongly suppressed when neighbors were present. These data indicated that flooding, salinity, and competition all played a role in determining the zonation pattern of S. salsa. Furthermore, the importance of salinity was found to vary with life-history stage. Based on the results from these field manipulative experiments, we suggest that the marsh plant zonation paradigm may hold true for plant distributions along landscape-scale topographic gradients from mudflats to uplands in general. The relative importance of flooding, salinity, and competition, however, may vary at different elevations within a site and between sites. Handling editor: Pierluigi Viaroli     

Blocking Phragmites australis reinvasion of restored marshes using plants selected from wild populations and tissue culture

This study tested a vegetation strategy for controlling Phragmites australis invasion into brackish marshes as an alternative to the current technique of repeated herbicide sprays followed by burning. This strategy involves blocking P. australis by planting desired plants selected from wild populations and/or tissue culture regenerants at key points on the major routes of P. australis invasion. The planting of native species was conducted at three sites in a herbicide-treated P. australis marsh near Salem, NJ. Wild population selections of three upland marsh shrubs, Myrica cerifera, Baccharis halimifolia, and Iva frutescens, as well as two grass species, Spartina alterniflora and Spartina patens, and two rushes, Juncus gerardi and Juncus roemerianus, were planted according to their normal zonation positions. Tissue culture regenerated plants of the two grasses and two rushes, and the sedge species Scirpus robustus, were also planted. Plant growth at each site was monitored each year after planting for up to 3 years. Most plants of B. halimifolia, I. frutescens, J. roemerianus, and S. patens demonstrated a consistent vigorous growth at all three sites, whether or not the plants were collected from wild populations or were tissue culture regenerants. These multi-layered walls of plants demonstrated effectiveness in controlling the P. australis by restricting or inhibiting its spread. Upon screening 48 regenerated plants of S. patens at one of the three sites, we found that some regenerants showed enhanced characteristics for blocking P. australis, such as greater expansion and a high stem density. The availability of the tissue culture-regenerated plants of the native marsh species makes it possible to select lines from local genotypes that have desirable characteristics for wetland restoration projects, such as blocking P. australis reinvasion.     

Crustacean metamorphosis: an omics perspective

We quantified the independent impacts of flooding salinity, flooding depth, and flooding frequency on the native species, Phragmites australis and Scirpus mariqueter, and on the invasive species Spartina alterniflora in the Yangtze River Estuary, China. Total biomass of all three species decreased significantly with increasing salinity, but S. alterniflora was less severely affected than P. australis and S. mariqueter. Elevated flooding depth significantly decreased their live aboveground biomass of P. australis and S. mariqueter, while S. alterniflora still had high live aboveground biomass and total biomass even at the highest flooding depth. These findings indicated that S. alterniflora was more tolerant to experimental conditions than the two native species, and an unavoidable suggestion is the expansion of this non-native species in relation to the native counterparts in future scenarios of increased sea-level and saltwater intrusion. Even so, environmental stresses might lead to significant decreases in total biomass and live aboveground biomass of all three species, which would potentially weaken their ability to trap sediments and accumulate organic matter. However, the relatively high belowground-to-aboveground biomass ratio indicated phenotypic plasticity in response to stressful environmental conditions, which suggest that marsh species can adapt to sea-level rise and maintain marsh elevation.     

ANATOMICAL AND METABOLIC RESPONSES TO WATERLOGGING AND SALINITY IN SPARTINA ALTERNIFLORA AND S. PATENS (POACEAE)

The effects of waterlogging and salinity (25 or 325 mol m ³ NaCl) stressors on the anatomy and metabolism of the marsh grasses 5. alterniflora Loisel. and S. patens Aiton (Muhl.) were investigated in a V factorial greenhouse experiment over 30 d. Waterlogging and salinity in combination resulted in anatomical and metabolic responses in both species. Waterlogging reduced soil redox potential and decreased root-specific gravity significantly in both species. The inadequacy of aerenchyma development under hypoxia to support aerobic root respiration in S. patens was indicated by significant increases in root alcohol dehydrogenase (ADH) activity of 1,752% and 420%, respectively, in the low and high salinity treatments. ADH activity was not increased significantly by flooding of S. alterniflora. Proline concentrations in roots and leaves were low at low salinities and increased significantly at high salinities in both species, but only under drained conditions. Decrease in leaf elongation by high salinity occurred in drained, but not flooded treatments in both species. Under flooded conditions, leaf elongation was significantly greater in S. alterniflora than S. patens. Greatest leaf elongation occurred in flooded low salinity S. alterniflora plants that had the least proline. Although both species are adapted to waterlogging and salinity, S. alterniflora appears to be more tolerant of reducing soil conditions and less responsive to higher salinity than S. patens.     

盐度和淹水对长江口潮滩盐沼植物碳储量的影响

盐生植物是盐沼有机碳储存的"临时库",也是土壤有机碳累积的主要来源,其碳储量大小对盐沼生态系统"碳汇"功能的发挥十分重要。以长江口潮滩本地种芦苇(Phragmites australis)和海三棱藨草(Scirpus mariqueter),及入侵种互花米草(Spartina alterniflora)为研究对象,采用单因素盆栽实验,模拟分析淹水盐度(0、5、10、15、25和35)、淹水深度(0、10、20、40、60cm和80cm)和淹水频率(每天、每3天、每7天、每10天和每15天)变化对各盐生植物地上、地下和总体碳储量大小的影响。研究结果表明,随着淹水盐度增加,芦苇、互花米草和海三棱藨草地上部分与总体碳储量均显著降低。土壤盐度可分别解释其地上部分碳储量变异的47.2%、66.5%和72.7%,与总体碳储量变异的34.7%、45.0%和62.0%。随着淹水深度增加,芦苇地上部分、总体碳储量和海三棱藨草地上部分碳储量均显著降低,其变异的68.6%、28.5%和71.1%可由淹水深度变化(10—80cm)解释。互花米草在80cm淹水深度下仍有较高的地上部分碳储量和总体碳储量。3种盐生植物碳储量对淹水频率变化的响应差异均不显著,所有处理地下部分碳储量差异也未达到显著水平。总体而言,互花米草对水盐胁迫的耐受性要强于本地种芦苇和海三棱藨草。尽管互花米草和芦苇具有相对较高的碳储量,但水盐胁迫对其碳储量的显著抑制作用不容忽视。海三棱藨草碳储量本就不高,输入土壤的有机碳量较为有限,海平面上升及盐水入侵等逆境胁迫会使其对盐沼"碳汇"贡献更加微弱。     

Seed flotation and germination of salt marsh plants: The effects of stratification, salinity, and/or inundation regime

We examined the effects of cold stratification and salinity on seed flotation of eight salt marsh species. Four of the eight species were tested for germination success under different stratification, salinity, and flooding conditions. Species were separated into two groups, four species received wet stratification and four dry stratification and fresh seeds of all species were tested for flotation and germination. Fresh seeds of seven out of eight species had flotation times independent of salinity, six of which had average flotation times of at least 50 d. Seeds of Spartina alterniflora and Spartina patens had the shortest flotation times, averaging 24 and 26 d, respectively. Following wet stratification, the flotation time of S. alterniflora seeds in higher salinity water (15 and 36 ppt) was reduced by over 75% and germination declined by more than 90%. Wet stratification reduced the flotation time of Distichlis spicata seeds in fresh water but increased seed germination from 2 to 16% in a fluctuating inundation regime. Fresh seeds of Iva frutescens and S. alternflora were capable of germination and therefore are non-dormant during dispersal. Fresh seeds of I. frutescens had similar germination to dry stratified seeds ranging 25-30%. Salinity reduced seed germination for all species except for S. alterniflora. A fluctuating inundation regime was important for seed germination of the low marsh species and for germination following cold stratification. The conditions that resulted in seeds sinking faster were similar to the conditions that resulted in higher germination for two of four species.     

EVALUATION OF PROLINE ACCUMULATION IN THE ADAPTATION OF DIVERSE SPECIES OF MARSH HALOPHYTES TO THE SALINE ENVIRONMENT

Proline accumulation by eight major species of salt marsh halophytes was examined under growth chamber and field conditions. When the plants were exposed to increasing salinities in the growth chamber, they accumulated proline after a threshold salinity had been reached. Three general patterns were apparent. Limonium carolinianum (Walt.) Britt. and Junius roemerianus Scheele began to accumulate proline at 0.25 m NaCl with accumulations up to 63.6 μmoles per gram fresh weight at higher salinities. The C₄ grasses, Spartina alterniflora Loisel., Spartina patens (Aiton) Muhl., and Distichlis spicata (L.) Greene, had threshold salinity levels around 0.5 m NaCl and accumulated proline to 27.4 μmoles per gram fresh weight. The succulents, Salicornia bigelovii Torr., Salicornia virginica L., and Borrichia frutescens (L.) DC, did not accumulate proline until very high salinities (0.7 m) were reached. Water stress imposed by polyethylene glycol instead of NaCl caused similar proline accumulation in the species studied, but to different extents. Field measurements of proline content and soil salinities correlated well with the findings from growth chamber experiments. Rates of proline accumulation and breakdown in L. carolinianum were sufficient for osmotic adjustment by the plant to the changes in interstitial salinity in the marsh. The significance of proline accumulation as an adaptation to the salt marsh environment was species specific. We suggest that proline accumulation is of considerable importance in L. carolinianum and J. roemerianus, important to the C₄ grasses at certain times and in certain locations in the marsh, and of little importance in the succulents.     

Whole-plant gas exchange responses of Spartina alterniflora (Poaceae) to a range of constant and transient salinities

A two-chamber-system was used to study whole-plant gas exchange responses of Spartina alterniflora to long-term and transient salinity treatments over the range of 5 to 40 ppt NaCl. Lower photosynthetic rates, leaf water vapor conductances, belowground respiration rates, and higher aboveground respiration rates in plants adapted to 40 ppt NaCl were observed. Area-specific leaf weight increased with salinity, although the salt content of leaf tissues did not. A reduced rate of gross photosynthesis and higher aboveground respiration rate in 40-ppt NaCl plants significantly lowered the net whole-plant CO₂ gain below that of 5-ppt NaCl plants, while the net CO₂ gain of 25-ppt NaCl plants was intermediate. Within 6 hr of increasing the salinity of 5- and 25-ppt NaCl plants by 20 and 15 ppt NaCl, S. alterniflora responded by reducing leaf water vapor conductance, which in turn reduced the photosynthetic rate. This response was reversed by returning the plants to their original salinity, which indicates that S. alterniflora adjusts water loss and gas exchange in response to transient salinity stress by regulating stomatal aperture. On the other hand, decreasing salinity of the growth media of plants cultured at 25 and 40 ppt NaCl had little or no effect on gas exchange characteristics. This suggests that S. alterniflora adapts to constant salinity through fixed, salinity-dependent structural modifications, such as stomatal density.     

Phenotypic plasticity of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and <Emphasis Type="Italic">Phragmites australis</Emphasis> in response to nitrogen addition and intraspecific competition

Phenotypic plasticity of the two salt marsh grasses Spartina alterniflora and Phragmites australis in salt marshes is crucial to their invasive ability, but the importance of phenotypic plasticity, nitrogen levels, and intraspecific competition to the success of the two species is unclear at present. Spartina alterniflora Loisel. is an extensively invasive species that has increased dramatically in distribution and abundance on the Chinese and European coasts, and has had considerable ecological impacts in the regions where it has established. Meanwhile, Phragmites australis Cav., a native salt marsh species on the east coast of China, has replaced the native S. alterniflora in many marshes along the Atlantic Coast of the US. This study determined the effects of nitrogen availability and culm density on the morphology, growth, and biomass allocation traits of Spartina alterniflora and Phragmites australis. A large number of morphological, growth, and biomass parameters were measured, and various derived values (culm: root ratio, specific leaf area, etc.) were calculated, along with an index of phenotypic plasticity. Nitrogen addition significantly affected growth performance and biomass allocation traits of Spartina alterniflora, and culm density significantly affected morphological characteristics in a negative way, especially for Spartina alterniflora. However, there were no significant interactions between nitrogen levels and culm density on the morphological parameters, growth performances parameters, and biomass allocation parameters of the two species. Spartina alterniflora appears to respond more strongly to nitrogen than to culm density and this pattern of phenotypic plasticity appears to offer an expedition for successful invasion and displacement of Phramites australias in China. The implication of this study is that, in response to the environmental changes that are increasing nitrogen levels, the range of Spartina alterniflora is expected to continue to expand on the east coast of China.     

Ecological processes shaping Central Patagonian salt marsh landscapes

Plant zonation is one of the most conspicuous ecological features of salt marshes worldwide. In this work we used a combination of field transplant and greenhouse experiments to evaluate the importance of interspecific interactions and physical stress in the determination of the major plant zonation patterns in Central Patagonian salt marshes. There, Spartina alterniflora dominates the low marsh, and Sarcocornia perennis the high marsh. We addressed two questions: (i) What prevents Spartina alterniflora from colonizing the Sarcocornia perennis‐dominated high marsh zone? and (ii) What prevents Sarcocornia perennis from colonizing the Spartina alterniflora‐dominated low marsh zone? Our experimental transplants combined with neighbour exclusion treatments showed that the presence of Sarcocornia perennis negatively affects Spartina alterniflora, preventing it from surviving and/or spreading. Complementary field transplant and greenhouse experiments showed that Sarcocornia perennis did not survive the frequent tidal submersion by approximately 1.5 m of turbid seawater in the Spartina alterniflora zone, but its survival was independent of the presence of Spartina neighbours, and of the strong soil anoxia as well. Our results suggest that Spartina alterniflora is excluded by Sarcocornia perennis towards the low marsh, where frequent and prolonged submersion limit the survival of the latter. We provide and discuss key baseline information to facilitate the future design of ecophysiological experiments designed to accurately identify the exact mechanisms acting in every situation.