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.     

A NEW NITROGEN-FIXING CYANOPHYTE-HEPATIC ASSOCIATION: NOSTOC AND PORELLA

Porella navicularis, a common leafy liverwort in western North America, was found to possess numerous epiphytic Nostoc colonies. Although the abundance of colonies was variable, the association was found consistently throughout a broad geographic range. Unlike cyanobacteria in other hepatic assocations, the Nostoc occurred as distinct colonies harbored in crevices and curled margins of leaves. Heterocyst frequency was 3–7%. Acetylene reduction activity was present in 85% of samples examined, with an average value of 53.5 nmol C₂H₄∙g dry wt^-1∙hr^-1 and a maximum of 316 nmol C₂H₄∙g dry wt^-1∙hr^-1. This suggests that bryophyte nitrogen-fixing associations may be more important than previously realized.     

GERMINATION AND SEEDLING RESPONSE OF ATLANTIC AND GULF COASTS POPULATIONS OF SPARTINA ALTERNIFLORA

Germination response to thermoperiod and seedling response to photoperiod-thermoperiod treatments and to uniform field conditions were compared for 12 populations of Spartina alterniflora Loisel. from along the Atlantic and Gulf coasts. Germination above 50 % was attained by seeds from all populations in 25–10, 30–15, and 35–20 C alternating diurnal thermoperiods following three months storage in estuarine water at 2–3 C. Except for Massachusetts, Connecticut, and Virginia, seedlings of populations from Ocracoke Island, North Carolina, and northward produced significantly more total biomass in the long- than in the short-day photoperiod in the 30–26 C thermoperiod. Seedling biomass of populations southward of Ocracoke Island, North Carolina, was not significantly affected by photoperiod in either the 18–14 or 30–26 C thermoperiods. Seedlings of all populations from Georgia and northward were significantly shorter and produced significantly more culms in the short- than in the long-day photoperiod in the 30–26 C thermoperiod. Seedlings from Mississippi and all populations from Virginia and northward had significantly lower shoot to root plus rhizome ratios under the short- than the long-day photoperiod in the 30–26 C thermoperiod. Flowering occurred only in populations from Ocracoke Island, North Carolina, northward in the 30–26 C thermoperiod. In a field study, flowering occurred in a north to south sequence and in all populations by the end of the second growing season. Controlled environment and field seedling studies indicated that southern populations flowered later, exhibited longer growing periods, and were less sensitive to photoperiod than northern populations. New England, Mid-Atlantic, South Atlantic, and Gulf coast populations differed in height, color, flowering time, length of growing period, and morphology through two growing seasons in the field study.     

EFFECT OF AN AMMONIUM NITRATE PULSE ON THE GROWTH AND ELEMENTAL COMPOSITION OF NATURAL STANDS OF SPARTINA ALTERNIFLORA AND JUNCUS ROEMERIANUS

A nitrogen (ammonium nitrate) pulse of 200 kg ha“¹ was added to stands of tall (1.0–1.5 m) Spartina alterniflora, short (< 0.5 m) Spartina alterniflora, and Juncus roemerianus in a Georgia salt marsh in July. The major response ten weeks later was an increase in the aerial biomass and a sharp reduction in the C/N ratio in short Spartina alterniflora. One year after the treatment the difference between the biomass in enriched and control plots was greater than ten weeks after treatment, but the C/N ratio in the plants in the treated plots had risen to that of the controls. The availability of nitrogen appears to limit growth in the middle elevation Georgia salt marsh (short S. alterniflora), but not in the lower (tall S. alterniflora) or higher (J. roemerianus) portions.     

EFFECTS OF O2 ON AMMONIUM UPTAKE AND ROOT RESPIRATION BY SPARTINA ALTERNIFLORA

Previously the growth of Spartina alterniflora has been found to be limited by nitrogen and correlated with sediment redox potential. In this study we have investigated a possible connection between these two factors. We have found that internal O₂ transport is insufficient to saturate NH₄⁺ uptake in short S. alterniflora in hydroponic culture. Rates of NH₄⁺ uptake and root respiration were very sensitive to O₂ concentration in the rhizosphere, saturating at about 5% O₂. Ammonium uptake continued at a reduced rate for at least 4 hr under anaerobic conditions. Plant to plant variations in anaerobic rates of NH₄⁺ uptake and root respiration were significantly correlated to the diffusion rate of CH₄ tracer gas from the leaves to the roots of individual plants.     

THE LIFE CYCLE OE SARGASSUM HORNERI (PHAEOPHYTA) IN LABORATORY CULTURE1

The life cycle of the large dioecious alga Sargassum horneri (Turner) C. Agardh was completed in unialgal culture by controlling photoperiod in relation to the phase of growth. Embryos isolated from a naturally grown female thallus gave rise to early germlings that rapidly formed blades under both short-day (9 h L) and long-day (15 h L) conditions at 20° C Shoot elongation, which followed early blade formation, occurred under the short-day conditions hut not under the long-day conditions. Functional female and male receptacles developed when thalli 8–14 cm long grown under the short-day conditions were transferred to the long-day conditions; gamete fusion occurred when male and female thalli were grown together. Fertilized oospores gave rise to normal thalli in a manner similar to that for in situ plants. Thus, the life cycle of S. horneri was completed in laboratory culture.     

Continuous fermentation to produce fungal protein. Effect of growth rate on the biomass yield and chemical composition of Fusarium moniliforme

Fusarium moniliforme was grown on a carob aqueous extract in a chemostat for fungal protein production. The substrate was adjusted to provide 0.5% carob sugars supplemented with inorganic salts. The dilution rate varied from 0.086 to 0.227 hr^?1 under constant conditions of temperature (30°C), pH (4.5), and oxygen saturation (60–80%). A yield of 0.709 g dry mycelium/g consumed carob sugar and a productivity value of 0.687 g dry mycelium/liter hr^?1 were obtained at μ = 0.205 hr^?1. The maintenance coefficient was 0.077 g carob sugar/g dry mycelium hr^?1. While the carbohydrate and purine content of dry mycelium increased at μ values from 0.114 to 0.205 hr^?1 both true (Lowry) and crude (N × 6.25) protein contents decreased at the same μ range. Maximum values of 36.3% true and 47.9% crude protein of dry mycelium were obtained at μ = 0.114 hr^?1, whereas a minimum purine content of 99.8 μmol/g corresponding to 6.42% nucleic acids was recorded at μ = 0.086 hr^?1. It was concluded that a continuous fermentation of carob aqueous extract using F. moniliforme should be operated at growth rates of approximately 0.205 hr^?1 in order to maximize protein production.     

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.     

TILLERING AND MORTALITY OF THE SALT MARSH CORDGRASS,SPARTINA ALTERNIFLORA

Recruitment and mortality of Spartina alterniflora tillers and culms were estimated from measurements of live standing crops at Wallops Island, Virginia. Stands of tall and medium form grass produced from 452 to 1,227 tillers m⁻² yr⁻¹. Those tillered in spring lengthened into culms their first year, while those tillered in late summer and fall may have overwintered and lengthened into culms their second year. Biennial growth was most prevalent in tall form Spartina. Fifty-six to sixty-one percent of all tillers died before lengthening into culms. Of the remainder, 9 to 56% died as culms before the time of flowering. Sixty-three percent of the surviving culms of tall form Spartina flowered, compared to only 14 to 23% in medium form. Mortality of tillers and culms removed 16 to 35% of the annual production of Spartina biomass from live standing crops before the time of flowering.     

14C UPTAKE BY THE MARINE ANGIOSPERM PHYLLOSPADIX SCOULERP

Rhizomes and attached leaves of Phyllospadix scouleri Hook, were collected in the intertidal zone along the central California coast and exposed to a solution of NaH¹⁴CO₃ in seawater under controlled laboratory conditions. Over a 90-min period roots and rhizomes absorbed very little ¹⁴C compared to leaves. Translocation during that time was minor. Plants pretreated with the photosynthetic inhibitor DCMU showed no ¹⁴C uptake, indicating that under normal circumstances the carbon which is absorbed by leaves is fixed and accumulates as photosynthate. The rate of gross photosynthesis was about 13 mg CO₂ g dry wt⁻¹ hr⁻¹. Gross photosynthesis of wet leaves exposed to ¹⁴CO₂ in air was significantly less than leaves exposed to NaH¹⁴CO₃. The effect of a leaf-grazing limpet (Notoacmea paleacea) on leaf anatomy and ¹⁴C uptake is discussed.     

THE EFFECT OF AERATION ON THE GROWTH OF SPARTINA ALTERNIFLORA LOISEL.

A greenhouse experiment was designed to investigate the correlations between waterlogging and aeration, and associated changes in pH, redox potentials and sulfide concentrations, on the growth of Spartina alterniflora Loisel. Elemental concentrations of the aerial and root material were determined and used for correlations with growth response. Redox potentials adjusted to pH 7 (Eh 7) ranged from −184 mv to 5 mv and were highly correlated (r) with aerial and root dry weight biomass (.97 and .97, respectively) and plant height (1.0). The range of soil pH at the conclusion of the study was 6.07 to 6.74 and was negatively correlated with aerial and root dry weight biomass. Sulfide concentrations ranged from 10–² to 10–⁷ m and vorrelations with aerial and root dry weights and height were −.85, −.85 and −.87, respectively. High negative correlations were found between sodium and sulfur concentrations and S. alterniflora growth. Positive correlations between potassium, phosphorus, manganese, zinc, copper, iron and growth response were also observed. Correlations of elemental concentrations of the plants with redox potentials and/or pH suggest that these two physical variables may be responsible in part for the regulation of 5. alterniflora growth in nature by regulating availability of nutritional elements.     

Ascomycete fungal communities associated with early decaying leaves of Spartina spp. from central California estuaries

Ascomycetous fungi play an important role in the early stages of decomposition of Spartina alterniflora, but their role in the decomposition of other Spartina species has not been investigated. Here we use fingerprint (terminal restriction fragment length polymorphism) and phylogenetic analyses of the 18S to 28S internal transcribed spacer region to compare the composition of the ascomycete fungal communities on early decay blades of Spartina species (Spartina alterniflora, Spartina densiflora, Spartina foliosa, and a hybrid (S. alterniflora × S. foliosa)) collected from three salt marshes in San Francisco Bay and one in Tomales Bay, California, USA. Phaeosphaeria spartinicola was found on all samples collected and was often dominant. Two other ascomycetes, Phaeosphaeria halima and Mycosphaerella sp. strain 2, were also common. These three species are the same ascomycetes previously identified as the dominant fungal decomposers on S. alterniflora on the east coast. Ascomycetes appeared to exhibit varying degrees of host specificity, demonstrated by grouping patterns on phylogenetic trees. Neither the exotic S. alterniflora nor the hybrid supported fungal flora different from that of the native S. foliosa. However, S. densiflora had a significantly different fungal community than the other species, and hosted at least two unique ascomycetes. Significant differences in the fungal decomposer communities were also detected within species (two clones of S. foliosa), but these were minor and may be due to morphological differences among the plants.     

Characterization of microsatellite loci in Spartina species (Poaceae)

The cordgrasses in the genus Spartina have become model organisms for studying biological invasions from both ecological and genetic perspectives. Here we characterize 11 disomic loci in Spartina alterniflora that show promise for population studies and for studying hybridization events between S. alterniflora and S. foliosa. Comparisons among invasive and native S. alterniflora populations showed that levels of allelic variation are lower in invasive populations. In addition, nearly all loci that amplified in S. foliosa populations and in a swarm of S. alterniflora×foliosa hybrids were polymorphic. We also found that several loci amplified successfully in other Spartina species.     

Induction of adult diapause and of low and high reproductive states in a parasitoid wasp,Ooencyrtus nezarae, by photoperiod and temperature

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) was reared on eggs ofRiptortus clavatus (Thunberg) (Heteroptera: Alydidae) at various temperatures under long-day (L16:D8) or short-day (L10:D14) conditions. There was no diapause during egg, larval or pupal stages under any set of conditions examined. However, at 15°C under short-day conditions, vitellogenesis was arrested in all adult females and they entered diapause. At 15°C under long-day conditions, or at 20°C under short-day conditions, some adult females entered diapause. Under the latter set of conditions, the adult females laid eggs but they laid fewer eggs than under long-day conditions, Even at 25°C, under short-day conditions, adult females laid fewer eggs than under long-day conditions, and this low rate of oviposition was attributed to the retarded development of ovaries. Diapause adults reared at 15°C were more resistant to low temperature than nondiapause adults reared at 25°C.     

Extent and degree of hybridization between exotic (Spartina alterniflora) and native (S. foliosa) cordgrass (Poaceae) in California, USA determined by random amplified polymorphic DNA (RAPDs)

Spartina alterniflora, smooth cordgrass, native to the eastern USA, was introduced into south San Francisco Bay ≈ 25 years ago. It has spread by purposeful introduction of rooted plants and dispersal of seeds on the tides. Previous work suggested that S. alterniflora was competitively superior to the native California cordgrass, S. foliosa, and that the two species hybridized. The present study determined the spread of S. alterniflora and S. foliosa × alterniflora hybrids in California and examined the degree of hybridization. We used nuclear DNA markers diagnostic for each species to detect the parental species and nine categories of hybrids. The California coast outside San Francisco Bay contained only the native species. All hybrid categories exist in the Bay, implying that several generations of crossing have occurred and that hybridization is bidirectional. Hybrids were found principally near sites of deliberate introduction of the exotic species. Where S. alterniflora was deliberately planted, we found approximately equal numbers of S. alterniflora and hybrid individuals; S. foliosa was virtually absent. Marshes colonized by water-dispersed seed contained the full gamut of phenotypes with intermediate-type hybrids predominating. The proliferation of hybrids could result in local extinction of S. foliosa. What is more, S. alterniflora has the ability to greatly modify the estuary ecosystem to the detriment of other native species and human uses of the Bay. To the extent that they share these engineering abilities, the proliferation of cordgrass hybrids could grossly alter the character of the San Francisco Bay.     

EFFECTS OF LIGHT AND HORMONES ON REGENERATION OF CRASSULA ARGENTEA FROM LEAVES

Plantlet regeneration from leaf segments of Crassula argentea was studied under variable light and hormone treatments. The effects of two hormones (indole 3-acetic acid and isopentenyl adenine at 10⁻⁴ to 10⁻¹⁰ m ) and photoperiods of 8 (short days) or 16 hr of light (long days) were examined. Exogenously applied hormones had little stimulating effect on regeneration but were consistently inhibitory at 10⁻⁴ m . Short days and continuous darkness caused the greatest promotion of regeneration. Prolonged short-day growth periods prior to culturing caused an increasingly greater regeneration response.     

Native and Introduced Ecosystem Engineers Produce Contrasting Effects on Estuarine Infaunal Communities

Cordgrasses in the genus Spartina are good examples of ecosystem engineers that modify habitat structure in estuaries throughout the world. In San Francisco Bay, California, USA, marshes containing native California cordgrass (Spartina foliosa) are being invaded by a hybrid (S. alterniflora × S. foliosa) formed after introduction of S. alterniflora. This study compared vegetation, sediment structure, and infaunal invertebrates in native and invaded marshes. We hypothesized that differences in the physical structure between S. foliosa and hybrid Spartina would be reflected in differences in density, biomass, diversity, and taxonomic composition of infauna. Hybrid Spartina modifies habitat structure more than S. foliosa by producing taller stems, and greater plant biomass both above- and belowground while occupying a much wider tidal range, thereby transforming open mudflats to a vegetated habitat. In general, S. foliosa areas contained significantly higher densities of benthic infauna than adjacent mudflats, while hybrid Spartina areas never contained greater infaunal densities than mudflats. This is because S. foliosa produces a moderate level of structure that can facilitate benthic invertebrates, whereas hybrid Spartina produces so much structure, particularly belowground, that it actually excludes invertebrates. Therefore, we suggest that these two closely related species both act as ecosystem engineers, but with opposing effects on invertebrate communities.     

Effects of salinity and temperature stress on Ecophysiological characteristics of exotic cordgrass, Spartina alterniflora

Salintiy and temperature are two important ecological factors which affect the distribution and abundance of Spartina alerniflora Loisel. To find out how cordgrass adapts to the environmental conditions in the introduced range, we studied the dynamics of a series of important physiological components including superoxide dismutase (SOD), guaiacol peroxidase (POD), catalase (CAT), malondialdehyde (MDA), soluble sugar and free proline under different salinity and temperature stresses. The results showed that low NaCl concentration (lower than 100 mmol·L⁻¹) enhanced the growth of S. alterniflora. On the contrary, high NaCl concentration (higher than 100 mmol·L⁻¹) inhibited the growth of S. alterniflora. To a certain extent, S. alterniflora was able to be acclimated to the osmotic pressure created by external solution concentration by adjusting the activities of POD, SOD and CAT, and the contents of free proline and soluble sugar. S. alterniflora varied in its responses to environment in different parts of the plant under 5°C and 38°C temperature stress. Compared with roots, leaves accumulated more soluble sugar, and CAT activities in leaves were higher, whereas SOD and POD activities in leaves were much lower than those in roots.     

Gibberellin A20 content of Bryophyllum daigremontianum under different photoperiodic conditions as determined by gas-liquid chromatography

Summary The gibberellin A₂₀ content of the long-short-day plant Bryophyllum daigremontianum (R. Hamet and Perr.) Berg. under different photoperiodic conditions was determined by gas-liquid chromatography. Purified extracts from ten plants were adequate for quantitative analysis by this method. The level of GA₂₀ increased following transfer from long-day (LD) to short-day (SD) conditions until after 38 SD it was three times higher on a dry weight basis than in comparable plants under continuous LD. No GA₂₀ could be detected in extracts of plants under permanent SD. These results are in agreement with earlier data obtained by assaying Bryophyllum extracts with the d-5 mutant of corn.     

DAYLENGTH AND LIGHT RESPONSES IN GROWTH AND FERTILITY OF GLOSSOPHORA KUNTHII (PHAEOPHYTA,DICTYOTALES) FROM PACIFIC SOUTH AMERICA1

Excised ligulae of Glossophora kunthii (C. Ag.) J. Ag. were cultured in photoperiods of 4–24 h and photon fluence rates of 10–75 μmol.m^?2.s^?1. Daylength interacted with irradiance on the growth of the ligulae. Maximal growth of primary ligulae occurred in long-day regimens with high irradiances suggesting an effect of irradiance on photosynthesis and growth. In contrast, growth of secondary ligulae was greatest in short-day regimes. Differences were significant at the highest irradiance tested. Differentiation of tetrasporangia on the ligulae is a short-day photoperiodic response. Daylengths of 8.5 h or less induced a sharp increase in numbers of fertile ligulae and tetrasporangia attaining maturity. Interruptions of the dark period decreased the development of tetrasporangia; the number of interruptions had a cumulative inhibitory effect. Differentiation of reproductive structures was influenced by interactions of photoperiod and irradiance. Maximum numbers of tetrasporangia were formed at short-day regimes and low irradiances; differentiation was completely inhibited at long-day conditions and high irradiance.