A Study on Polymorphism in Aster tripolium L. (Sea Aster)

Abstract: Genotypic and environmental variation in Aster tripolium L. was studied in common garden experiments and in transplantation experiments in different saltmarsh sites along the estuarine gradient of the Westerschelde and along the elevation gradient of individual saltmarshes. Analysis of the variation in a number of morphologic characters of the inflorescences indicated that this variation is both environmentally as well as genetically controlled. Morphologic differences between an Aster tripolium population of a brackish marsh and a number of populations of a saline marsh were unchanged whether the plants were growing in their original habitat or in a transplant habitat. The population from the brackish marsh had smaller inflorescences with ray florets and a predominantly biennial life cycle, in contrast with the populations of the saline marsh, which had larger inflorescences usually without or with a few ray florets and showed a tendency to a perennial life cycle. We concluded that the variation between the populations of the saline and the brackish marsh is mainly genetically controlled, while the variation within the saline marsh is mainly environmentally controlled.     

Salt tolerance in Aster tripolium L. II. Ionic regulation

Abstract Measurements of tissue ion contents (Na, K and Cl) were carried out at frequent intervals on plants of Aster tripolium L. grown at a range of salinities for 36 d. Aster tripolium behaved as a typical halophyte showing high levels of inorganic ion accumulation even at low salinities. As salinity increased Na replaced K to a large extent in the shoot but root K was unaffected up to 500 mol m^?3 external NaCl. Shoot (Na + K) concentration on a tissue water basis was maintained constant in all treatments throughout the experiment, whereas shoot (Na + K) on a dry weight basis showed marked fluctuations in some treatments. An increase in (Na + K) per gram dry weight was, however, accompanied by a parallel increase in fresh weight: dry weight (FW : DW) ratio. Transport of (Na + K) to the shoot per unit root weight changed during the experiment in the manner expected, given the observed changes in shoot relative growth rate and FW : DW to result in a constant shoot (Na + K) concentration on a water basis. Chloride was the major balancing anion in the shoot at high salinity, but never accounted for more than 38% of the (Na + K) found in the root tissue. At all salinities (Na + K) salts accounted for the majority of the measured shoot sap osmotic potential. The interactions between salinity, growth, ion transport and osmotic adjustment are discussed.     

Salt tolerance in Aster tripolium L. I. The effect of salinity on growth

Abstract A study of the growth of the maritime halophyte Aster tripolium L. has been carried out over a range of salinity treatments. The regression approach to growth analysis using frequent small harvests has been used to allow ‘continuous’ measurement of growth over a period of 36 d. Salinity was applied with the major ions present in ratios typical of those found in seawater. Growth was inhibited in terms of both dry weight production and leaf expansion at salinity levels equivalent to 0.625 strength sea water (full culture solution 300) and above, with the greatest effect being seen in terms of leaf area. Aster tripolium did not show increased succulence at high salinity, leaf fresh weight to dry weight ratio in fact declined, whilst leaf fresh weight per unit area remained constant. It should be noted that the plants exhibit low growth rates due to the low light intensity used.     

Population Variability of Aster tripolium L. of Zhoushan Islands as Related to Habitat

In the identification of the specimens collected from Zhoushan (Chushan) Islands of Zhejiang (Chekiang) Province, the authors have found that the species Aster tripolium L. (Tripolium vulgate Nees) of coastal salt marshes has a rather high amplitude of morphological variation. We, then, made a field investigation in Zhoushan Islands in October of 1980. A mass collection of five different populations of the same species of different ecological sites was carried out. Quantitative measurements of totally eight morphological characters, namely, the height of individual plants, the length and breadth of leaves and ligulate flowers, the number of lower branches, the degree of promineney of tap-roots, and the color of the base of stems, were made. The resulting data of these measurements and calculations were then compared by using methods of E. Anderson's pietorialized scatter diagram and standard deviation in order to bring to light the differentiation pattern of that species occurred in response to different habitats. Samples of soil of different population sites were also collected; their pH values and contents of NaCl were examined. According to the variation pattern of phenotypes as shown in these diagrams and figures, the populations of Aster tripolium of Zhoushan Islands could be roughly divided into the following two types: (1) the tall and long-leaved type of lower marshes with a soil salinity of 0.54%–0.56% of NaCl, including populations Nos. 1 and 2, with plants 49–51 cm. high (mean value), very few or no lower branches, usually dark purplish at the base of stems, and often without prominent tap-roots; (2) the low and short-leaved type of higher marshes and saline seepage areas above the limits of tidal submergence, with a soil salinity of 0.06%–0.17% of NaC1, including populations Nos. 3 and 4, with plants 19–26 cm. high (mean value), a few to manylower branches, and usually greyish white or slightly purplish at the base of stems. It is worth of note that these two different types of phenotypic variation of populations are on the whole corresponding to the two habitat types, which differ from each other chiefly in soil salinity resulted from the different levels of the sites, particularly the distance between the population sites and the salterns, and in the density of plants as well, which could probably be related to the intensity of competition. It is also interesting to note that owing to the site of population No. 5 being subjected to trampling, grazing and moving, the plants are usually mechanically stripped of their terminal buds. This is probably why the members of that population often show a peculiar pseudo- prostrate habit with their lower branches sometimes spreading at right angles to the main stem. However, the pattern of the population differinces obtained is often predetermined by the pattern of sampling. The real pattern of population differentiation of Aster tripolium is more likely clinal instead of abrupt. Without comparative experimental cultivation, it is difficult to decide whether the phenotypic variations mentioned above are genetically fixed or simply environmental modifications. Nevertheless, A. J, Gray et al. (1979) has shown that the large differences exist between populations of Aster tripolium at the different levels on the British salt marshes appear in uniform cultivation and may have a large genetic component. So, it might be expected that both of the factors-genetical and environmental, are likely to play their parts in the population differentiation of the said species. According to the classification of population systems given by V. Grant (1963), the two different types of populations of Aster tripolium of Zhoushan Islands may be considered as two ecological races.     

Elevated atmospheric CO2 concentration enhances salinity tolerance in Aster tripolium L.

Our study aimed at investigating the influence of elevated atmospheric CO₂ concentration on the salinity tolerance of the cash crop halophyte Aster tripolium L., thereby focussing on protein expression and enzyme activities. The plants were grown in hydroponics using a nutrient solution with or without addition of NaCl (75% seawater salinity), under ambient (380 ppm) and elevated (520 ppm) CO₂. Under ambient CO₂ concentration enhanced expressions and activities of the antioxidant enzymes superoxide dismutase, ascorbate peroxidase, and glutathione-S-transferase in the salt-treatments were recorded as a reaction to oxidative stress. Elevated CO₂ led to significantly higher enzyme expressions and activities in the salt-treatments, so that reactive oxygen species could be detoxified more effectively. Furthermore, the expression of a protective heat shock protein (class 20) increased under salinity and was even further enhanced under elevated CO₂ concentration. Additional energy had to be provided for the mechanisms mentioned above, which was indicated by the increased expression of a β ATPase subunit and higher v-, p- and f-ATPase activities under salinity. The higher ATPase expression and activities also enable a more efficient ion transport and compartmentation for the maintenance of ion homeostasis. We conclude that elevated CO₂ concentration is able to improve the survival of A. tripolium under salinity because more energy is provided for the synthesis and enhanced activity of enzymes and proteins which enable a more efficient ROS detoxification and ion compartmentation/transport.     

Responses of the stomata of Aster tripolium to calcium and sodium ions in relation to salinity tolerance

In previous work, the stomata of the maritime halophyte Astertripolium L. were shown to close when NaCl concentrations risein the vicinity of the guard cells. Further studies have nowrevealed important effects of calcium on the ionic responsesof the stomata. When the guard cells were presented with KCl,Ca²⁺ suppressed opening in a manner similar to that which hasbecome familiar in other species such as Commelina communisL. However, in the presence of NaCl, Ca²⁺ had the opposite effect,reducing the closing response to NaCl. This pattern of behaviouris discussed in relation to known salt effects on membranes,but the underlying physiological basis remains obscure. A previous study led to the hypothesis that the closing responseof the stomata to Na⁺ ions may make an important contributionto the salinity tolerance of this species. Here we report thatincreasing supplies of Ca²⁺ ions reduce the effect of salinityon stomatal conductance in the whole plant as well as in theisolated epidermis. This finding is consistent with the wellestablished role of calcium in increasing resistance to salinity:in the presence of high calcium the plant can tolerate a greatersalt intake, and hence there is a reduced need for transpirationto be restricted by partial stomatal closure. Key words: Sodium, calcium, Aster tripolium, stomata, salinity tolerance     

Aster tripolium L. and Sesuvium portulacastrum L.: two halophytes, two strategies to survive in saline habitats.

Aster tripolium L. (Dollart, Germany) and Sesuvium portulacastrum L. (Dakhla, Morocco) are potential halophytic vegetables, fodder plants, and ornamentals for re-vegetating saline land. To compare their strategies involved in salt tolerance both plants were grown with 0%, 1.5%, and 3% (Aster) or 0%, 2.5%, and 5% (Sesuvium) NaCl in the watering solution. The growth rate was reduced in both species with increasing NaCl concentrations. The quotient of Na(+)/K(+) indicates that Aster accumulates more K(+) in comparison to Na(+) while the reverse is true for Sesuvium. Osmolality of the leaf sap increased with increasing NaCl concentration in both Aster and Sesuvium. Transpiration rate was severely reduced in both Aster (3%) and Sesuvium (5%) plants after 10 d of NaCl watering. The CO(2) assimilation rate decreased in Aster (3%) and Sesuvium (5%) NaCl-treated plants from day 5 to day 10. The most important results from chlorophyll fluorescence measurements were derived from the non-photochemical quenching analysis (NPQ). First, both plants had linearly increasing levels of NPQ with increasing NaCl concentrations. Second, Sesuvium had almost half the NPQ value when compared to Aster under increased soil salinity. In Aster P-ATPase activities were decreased in plants treated with 3% NaCl after three days of treatment, F-ATPase activities increased with increasing NaCl concentrations and no clear changes were measured in V-ATPase activities. In Sesuvium any changes could be observed in the three ATPase activities determined. To conclude, Aster and Sesuvium use different strategies in adaptation to soil salinity.     

Salt tolerance in Aster tripolium L. III. Na and K fluxes in intact seedlings

Abstract Uptake and transport of Na and K was studied using the radioactive tracers ²²Na and ⁴²K in intact Aster tripolium L. seedlings grown at two salinities CS 10 and CS 100, (containing 10mol m^?1 and 100 mol m^?3 Na, respectively, together with other major ions in the proportions found in sea water). At both salinities a much greater proportion of the Na than K taken up by the plant was subsequently transported to the shoot. Most ⁴²K fluxes were reduced by about 40% in CS 100 plants relative to CS 10 except root accumulation which increased. Experiments involving changing the salinity from CS 10 to CS 100 showed that ⁴²K fluxes remained constant for at least 40 h, indicating that competition with Na for uptake sites was not the cause of the reduced flux in CS 100 plants. ²²Na fluxes responded immediately to a change in salinity with all fluxes increasing six-fold when the salinity was raised. When the salinity was lowered, however, root accumulation returned to the level in CS 10 control plants whereas transport to the shoot was inhibited by the previous high salinity treatment, being reduced to only 35% of the rate in CS 10 plants. The time courses of osmotic adjustment and Na accumulation following an increase in salinity were found to be very similar, with sufficient Na being accumulated to account for the observed increase in sap osmotic pressure.     

Nitrate accumulation and growth of Aster tripolium L. with a continuous and intermittent nitrogen supply*

Abstract The ‘tidal salt marsh’ ecotype of the halophyte Aster tripolium L. was grown in a nutrient solution with either a continuous or an intermittent NO₃^? supply with either Cl^? or SO₄^2? as the alternative anion. With increasing periods of NO₃^? supply per week the rate of the dry weight increment increased. When NO₃^? was supplied for longer than 48 h per week, the dry weight and the organic-N content in the shoots hardly increased, whereas the NO₃^? content in shoots and roots increased further. With alternated supply of a nutrient solution containing NO₃^? with one containing Cl^?, the internal NO₃^? content in the shoot was lower than in shoots grown in solutions in which NO₃^? alternated with SO₄^2?. It is concluded, that NO₃^? does not have a specific function in osmoregulation.     

Growth and protein profile responses in the halophyte sea aster (Aster tripolium L.) suspension-cultured cells to salinity

We established salt tolerant cell lines, which survived and grew under high salinity conditions with 150 mM (S-150) and 300 mM (S-300) NaCl, to study the effects of salt stress on the proliferation and protein profile of these cells in the halophyte sea aster,Aster tripolium L. These salt-adapted cell lines were produced from leaves and selected by repeated suspension subculture in media containing NaCl every 25 days for five cycles. S-150 cells displayed no inhibition in their growth compared to control cells maintained under non-stressed conditions. S-150 cells exhibited approximately a 15-fold increase in both fresh and dry weight during the 25 days under saline conditions. S-300 cells showed positive growth under severe salt stress, but their dry matter gain was significantly less than that of the S-150 cells, with only a 2.5-fold increase in dry weight. We also detected changes in the protein profile of salt-adapted cells with two specifically induced polypeptides (basic 58.4 and acidic 24.8 kDa) and one enhanced polypeptide (basic 15.1 kDa) in the soluble fraction, and one specifically induced polypeptide (42.0 kDa) in the insoluble fraction.     

Arbuscular mycorrhizal fungi enhance root cadmium and copper accumulation in the roots of the salt marsh plant Aster tripolium L.

It is known that vegetation plays an important role in the retention of heavy metals in salt marshes by taking up and accumulating the metals. In this study, we investigated whether arbuscular mycorrhizal fungi (AMF) increase Cd and Cu uptake and accumulation in the root system of the salt marsh species Aster tripolium L., and whether indigenous AMF isolated from polluted salt marshes have higher capacity to resist and alleviate metal stress in A. tripolium than isolates of the same species originated from non-polluted sites. Plants inoculated with Glomus geosporum, either isolated from a polluted salt marsh site (PL isolate) or from a non-polluted site (NP isolate), and non-mycorrhizal (NM) plants were compared in a pot experiment at four different Cd and Cu concentrations. Cd had no effect in root colonization, whereas high concentrations of Cu decreased colonization level in plants inoculated with the NP isolate. AM colonization did not increase plant dry weight or P concentration but influenced root Cd and Cu concentrations. Inoculation with PL and NP isolates enhanced root Cd and Cu concentrations, especially at highest metal addition levels, as compared to NM plants, without increasing shoot Cd and Cu concentrations. There was no evidence of intraspecific variation in the effects between AMF isolated from polluted and non-polluted sites, since there were no differences between plants inoculated with PL or NP isolate in any of the tested plant variables. The results of this study showed that AMF enhance metal accumulation in the root system of A. tripolium, suggesting a contribution of AMF to the sink of metals within vegetation in the salt marshes.     

Disentangling the photochemical salinity tolerance in Aster tripolium L.: connecting biophysical traits with changes in fatty acid composition

• A profound analysis of A. tripolium photochemical traits under salinity exposure is lacking in the literature, with very few references focusing on its fatty acid profile role in photophysiology.
• To address this, the deep photochemical processes were evaluated by Pulse Amplitude Modulated (PAM) Fluorometry coupled with a discrimination of its leaf fatty acid profile.
• Plants exposed to 125–250 mm NaCl showed higher photochemical light harvesting efficiencies and lower energy dissipation rates. under higher NaCl exposure, there is evident damage of the oxygen evolving complexes (OECs). On the other hand, Reaction Centre (RC) closure net rate and density increased, improving the energy fluxes entering the PS II, in spite of the high amounts of energy dissipated and the loss of PS II antennae connectivity. Energy dissipation was mainly achieved through the auroxanthin pathway. Total fatty acid content displayed a similar trend, being also higher under 125–250 mm NaCl with high levels of omega‐3 and omega‐6 fatty acids. The increase in oleic acid and palmitic acid allows the maintenance of the good functioning of the PS II. Also relevant was the high concentration of chloroplastic C16:1t in the individuals subjected to 125–250 mm NaCl, related with a higher electron transport activity and with the organization of the Light Harvesting Complexes (LHC) and thus reducing the activation of energy dissipation mechanisms.
• All these new insights shed some light not only on the photophysiology of this potential cash‐crop, but also highlight its important saline agriculture applications of this species as forage and potential source of essential fatty acids.

     

Elevated atmospheric CO2 concentration ameliorates effects of NaCl salinity on photosynthesis and leaf structure of Aster tripolium L.

This study investigated the interaction of NaCl-salinity andelevated atmospheric CO₂ concentration on gas exchange, leafpigment composition, and leaf ultrastructure of the potentialcash crop halophyte Aster tripolium. The plants were irrigatedwith five different salinity levels (0, 25, 50, 75, 100% seawatersalinity) under ambient and elevated (520 ppm) CO₂. Under salineconditions (ambient CO₂) stomatal and mesophyll resistance increased,leading to a significant decrease in photosynthesis and wateruse efficiency (WUE) and to an increase in oxidative stress.The latter was indicated by dilations of the thylakoid membranesand an increase in superoxide dismutase (SOD) activity. Oxidativestress could be counteracted by thicker epidermal cell wallsof the leaves, a thicker cuticle, a reduced chlorophyll content,an increase in the chlorophyll a/b ratio and a transient declineof the photosynthetic efficiency. Elevated CO₂ led to a significantincrease in photosynthesis and WUE. The improved water and energysupply was used to increase the investment in mechanisms reducingwater loss and oxidative stress (thicker cell walls and cuticles,a higher chlorophyll and carotenoid content, higher SOD activity),resulting in more intact thylakoids. As these mechanisms canimprove survival under salinity, A. tripolium seems to be apromising cash crop halophyte which can help in desalinizingand reclaiming degraded land. Key words: Aster tripolium, cash crop halophyte, elevated CO₂, gas exchange, oxidative stress, photosynthesis, salt tolerance, ultrastructure, water use efficiency Received 29 July 2008; Revised 8 October 2008 Accepted 9 October 2008     

A vegetation zonation from saltmarsh to riverbank in New Zealand

Abstract. Brackish riverbank marshes have been little studied. Therefore, a plant community sequence was sampled from saltmarsh to near-freshwater riverbank marsh on a number of disjunct marshes along the Taieri River, Otago, New Zealand, from near the mouth to 9 km inland. Salinity decreased steadily upstream, though the actual values were very different on two days sampled. Ten communities are recognised. The major vegetation zonation was upshore more than upstream, though there were several interactions between the upshore and upstream gradients. Few species, if any, were restricted to the mid reaches of the length of river sampled. There was only a very slight upshore increase in species richness, and no trend upstream. Sequences of communities occurred upshore on all marshes, but the sequence differed, even within a marsh. Species were assorted into communities in different ways from those of marine marshes in the area. Some species, native and exotic, occupied different beta-niches from those they occupied in other countries. Individualistic community structure is inferred.     

Genetic variation among different populations of Aster tripolium grown on naturally and anthropogenic salt-contaminated habitats: implications for conservation strategies

The sea aster, Aster tripolium L., grows naturally in temperate regions, mainly in the salt meadows close to the coast. The species is also found in naturally and anthropogenically salt-contaminated inland habitats, such as potash mine dumps. The genetic relationships among populations from different habitats and correlations of the genotype with physiological and vegetational parameters were investigated. A. tripolium plants from five different sites close to the seashore on the North Sea island Baltrum, from five different potash mine dumps and, as an outgroup, from the seashore in Japan were probed. DNA was extracted from five plants from each of the 11 A. tripolium populations and analyzed for random amplified polymorphic DNA (RAPD). Altogether 35 polymorphic bands in 51 individuals and 45 different detectable genotypes could be identified. For evaluation of the genetic variation using RAPD bands, the neighbor-joining method, the principal coordinate analysis, and the analysis of molecular variance were applied, resulting in the classification into three genetic groups. A. tripolium plants from different ecological habitats on Baltrum were closely related while the plants growing at the deposit dumps showed a higher genetic diversity. The Japanese population was genetically very different from the German populations. Correlations between phytosociological and soil parameters and the respective genotype were not significant. The results argue for a conservation of anthropogenically salt-contaminated habitats to maintain genetic variability not only on the species level, but also within a species.     

Avoidance of sodium accumulation by the stomatal guard cells of the halophyte Aster tripolium

X-ray microanalysis has revealed that the sodium content ofthe stomatal guard cells of Aster tripolium remains much lowerthan that of other leaf cells when the plants are grown at highsalinity. Large amounts of sodium did, in contrast, accumulatein epidermal and subsidiary cells, and particularly in the mesophylltissue, suggesting that a mechanism exists to limit the extentof its entry into guard cells. Even in plants grown at highsalinity, the content of potassium was much higher than thatof sodium in the guard cells, consistent with the view thatthis is a major ion involved in determining stomatal movementsin this halophyte. Determinations were also made for the nonhalophyte Commelinacommunis, and it was found that the guard cells accumulatedlarge amounts of sodium when it was presented to them as analternative to potassium. It is suggested that the acquisition by the guard cells of someability to restrict the intake of sodium ions may be an importantcomponent of sodium-driven regulation of transpiration, andhence of salinity tolerance, in A. tripolium. Key words: Salinity tolerance, sodium, potassium, stomata, Aster tripolium     

Photosynthetic limitations of a halophyte sea aster (Aster tripolium L) under water stress and NaCl stress

To understand the mechanisms of salt tolerance in a halophyte, sea aster ( Aster tripolium L.), we studied the changes of water relation and the factors of photosynthetic limitation under water stress and 300 mM NaCl stress. The contents of Na(+) and Cl(-) were highest in NaCl-stressed leaves. Leaf osmotic potentials ( Psi(s)) were decreased by both stress treatments, whereas leaf turgor pressure ( Psi(t)) was maintained under NaCl stress. Decrease in Psi(s) without any loss of Psi(t) accounted for osmotic adjustment using Na(+) and Cl(-) accumulated under NaCl stress. Stress treatments affected photosynthesis, and stomatal limitation was higher under water stress than under NaCl stress. Additionally, maximum CO(2) fixation rate and O(2) evolution rate decreased only under water stress, indicating irreversible damage to photosynthetic systems, mainly by dehydration. Water stress severely affected the water relation and photosynthetic capacity. On the other hand, turgid leaves under NaCl stress have dehydration tolerance due to maintenance of Psi(t) and photosynthetic activity. These results show that sea aster might not suffer from tissue dehydration in highly salinized environments. We conclude that the adaptation of sea aster to salinity may be accomplished by osmotic adjustment using accumulated Na(+) and Cl(-), and that this plant has typical halophyte characteristics, but not drought tolerance.