Salt resistance and the environment of the cell wall of some halophytes

It was shown for Cochlearia anglica roots in comparison with Phaseolus vulgaris, roots that: (a) the cell walls of the halophyte are heavy; (b) they have a high Ca²⁺ content; (c) they may be a convenient means of transfer for Na⁺.In Aster tripolium as well as in Atriplex littoralis NaCl alters the composition of the various pectic fractions and maintains a low pectinemethylesterase activity and a great cellulase activity. This effect may be responsible for the great plasticity of the cell walls which allows an increased cell growth and leads to the formation of succulent organs, therefore to the dilution of salt within the cells.     

A new species of Pemphigus Hartig (Hornoptera: Aphidoidea) from Western Europe

A new aphid species, Pemphigus trehernei, found on the roots of Aster tripolium L. in Western Europe is described. The gall, fundatrix and alate fundatrigenia were produced artificially on Populus nigra L. var. italica Duroi. The life cycle and host range of the species were investigated.     

The effect of lead contamination of soils and air on its accumulation in pollen

Summary Concentrations of heavy metals in the honey, derived fromAster tripolium was the starting-point for this investigation. Lead content of pollen inAster tripolium andTaraxacum officinale is mainly due to airborne lead. Accumulation of other heavy metals such as zinc and copper in pollen occur mainly via a translocation process from roots to flowers.     

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.     

Peculiarities of muscles and skeleton structure in <Emphasis Type="Italic">Campiglossa plantaginis</Emphasis> (Haliday, 1833) ovipositor (Diptera,Terthritidae) in association with larval mode of life

Sclerites and muscles of the ovipositor in the tephritid fly Campiglossa plantaginis (Haliday, 1833), with phytophagous larvae developing in inflorescences of Aster tripolium L. and other Asteraceae, were examined. Structural adaptations of the ovipositor, providing oviposition into different substrates, are discussed in relation to different trophic specialization of larvae.     

Stomatal responses to sodium ions in Aster tripolium: a new hypothesis to explain salinity regulation in above-ground tissues

A study has been made of the ionic relations of stomata of Aster tripolium L., a maritime halophyte which colonizes coastal saltmarshes. The results obtained allow us to add this species to the growing list for which an involvement of K⁺ transport in stomatal movements has been demonstrated. However, an additional and ecologically important characteristic was found: there was a suppression of stomatal opening by increasing NaCl concentrations. A new hypothesis is offered of the mechanism for controlling salt and water relations in A. tripolium, a species which does not possess glands or other means of excreting salt. It is suggested that when the capacity of the tissues to accumulate salt in cell vacuoles is exceeded, the concentration of Na⁺ ions in the apoplast around the guard cells begins to rise. This causes partial stomatal closure, reduces transpiration and increases water-use-efficiency. Therefore, the flow of salt into the leaves is reduced but growth (and the manufacture of the new photosynthates required to support it) can continue. Aster tripolium can be added to the small list of known species which readily yield isolated epidermis suitable for detailed stomatal studies. Throughout this study, we have compared its stomatal physiology with C. communis, which has been thoroughly investigated in the past.     

The performance of the leaf mining microlepidopteran Bucculatrix maritima (Stt.) on the salt marsh halophyte,Aster tripolium (L.), exposed to different salinity conditions

Summary The performance of phytophagous insects is influenced by the nutritional quality of the food plant, which may vary with environmental conditions. Hardly any information exists on food-plant mediated effects of variable soil salinity on the performance of phytophagous insects. Conspicuous differences in salinity levels, however, are found in soils of intertidal wetlands such as salt marshes and mangroves. The growth of larvae of Bucculatrix maritima, a leaf miner of the salt marsh halophyte Aster tripolium, was studied on the host plant along the salinity gradient of the Westerschelde estuary (S.W. Netherlands). In addition, its performance on A. tripolium grown on low or high salinity culture medium was investigated experimentally. Although salinity conditions significantly influenced the chemistry of the host plants, insect performance seemed almost unaffected, although near the mouth of the estuary high environmental salinities may have caused some inhibition of larval growth. The results contrast with our previous studies on the stem-borer Agapanthia villosoviridescens, which showed that growth and development was conspicuously influenced by the changing characteristics of Aster tripolium along the estuarine salinity gradient. The location-dependent qualities of halophytes in an estuary thus appear to have species-specific effects on insect performance. We hypothesize that this phenomenon contributes to the existence of non-identical distribution patterns of phytophagous insects associated with the same halophyte in an estuary.     

How do Mycorrhizas Affect C and N Relationships in Flooded Aster tripolium Plants?

The mycorrhizal associations established between plants and fungi have multiple effects on plant growth, directly affecting stress tolerance. This work aimed to explore arbuscular mycorrhizal (AM) effects on carbon and nitrogen relationships of Aster tripolium L. and consequently on its flooding tolerance. Mycorrhizal and non-mycorrhizal juvenile plants were submitted to non-flooding and tidal flooding conditions for 56 d. Tidal flooding reduced biomass, but the presence of mycorrhiza had an ameliorating effect. The AM symbioses seem to have, like flooding, a stressful effect on A. tripolium at an early stage of plant development. However, once the plant was established, an improvement of growth performance of plants with mycorrhiza under flooding conditions was observed. The better tolerance of AM plants to flooding was mediated through an improvement of the osmotic adjustment of the plant tissues (higher concentrations of soluble sugars and proline) and through the increment of nitrogen acquisition in tidal-flooded plants.     

High occurrence of rare inland halophytes on post‐mining sites in western Ukraine

Rare inland halophytic vegetation including the associations Salicornietum prostratae, Spergulario marginatae‐Suaedetum prostratae, Puccinellietum limosae, com. Puccinellietum limosae – a variant of Aster tripolium subsp. pannonicus and Astero pannonici‐Bolboschoenetum compacti have been found on 3 localities in the Drohobych region of western Ukraine. These are isolated (‘island’) localities, almost unknown until today. Halophytes grew there on salted sites in the vicinity of the old salt industry. Species of halophytes previously not known from this region were discovered: Aster tripolium subsp. pannonicus, Bolboschoenus maritimus, Gypsophila paniculata, Limonium gmelinii, Lotus tenuis, Puccinellia distans, Sonchus palustris, Suaeda prostrata, Trifolium fragiferum var. bonannii, Triglochin maritima, Typha laxmannii and Zannichellia palustris subsp. pedicellata. The occurence of these plants strictly depends on the presence of salt in the soil and they are endangered by shrinkage of salted sites following the termination of the salt industry in Drohobych region.     

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.     

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.     

Guard cell cation channels are involved in Na+–induced stomatal closure in a halophyte

The halophyte Aster tripolium, unlike well-studied non-halophytic species, partially closes its stomata in response to high Na⁺ concentrations. Since A. tripolium possesses no specific morphological adaptation to salinity, this stomatal response, preventing excessive accumulation of Na⁺ within the shoot via control of the transpiration rate, is probably a principal feature of its salt tolerance within the shoot. The ionic basis of the stomatal response to Na⁺ was studied in guard cell protoplasts from A. tripolium and from a non-halophytic relative, Aster amellus, which exhibits classical stomatal opening on Na⁺. Patch-clamp studies revealed that plasma membrane K⁺ channels (inward and outward rectifiers) of the halophytic and the non-halophytic species are highly selective for K⁺ against Na⁺, and are very similar with respect to unitary conductance and direct sensitivity to Na⁺. On the other hand, both species possess a significant permeability to Na⁺ through non-rectifying cation channels activated by low (physiological) external Ca²⁺ concentrations. Finally, it appeared that the differential stomatal response between the two species is achieved, at least in part, by a Na⁺-sensing system in the halophyte which downregulates K⁺ uptake. Thus, increases in guard cell cytosolic Na⁺ concentration in A. tripolium but not in A. amellus, lead to a delayed (20–30 min) and dramatic deactivation of the K⁺ inward rectifier. This deactivation is probably mediated by an increase in cytosolic Ca²⁺ since buffering it abolishes the response. The possible role of K⁺ inward rectifiers in the response of A. tripolium’s stomata to Na⁺, suggested by patch-clamp studies, was confirmed by experiments demonstrating that specific blockade of inward rectifying channels mimics Na⁺ effects on stomatal aperture, and renders aperture refractory to Na⁺.     

Impact of different sheep grazing intensities on salt marsh vegetation in northern Germany

Abstract. Mainland salt marshes in Schleswig-Holstein (northern Germany) have been grazed intensively by sheep for several decades. In 1988 experimental sites were established in the lower and middle salt marsh of Sönke-Nissen-Koog and subjected to different grazing intensities. From 1989 to 1993 the impact of sheep grazing on the composition and structure of the salt marsh vegetation was studied through the yearly analysis of permanent plots, vegetation mapping and measurements of the vegetation height. The intensively grazed site (10 sheep/ha) is covered by a short monotonous Puccinellia maritima sward with Salicornia europaea and Suaeda maritima. Halimione portulacoides and Aster tripolium, especially flowering plants, are rare. On the sites with 1.5 and 3 sheep/ha Puccinellia maritima remained dominant. The population density of Salicornia europaea decreases after reduction of the grazing intensity, whereas Suaeda maritima finds optimal growing conditions. Stands of Halimione portulacoides and flowering Aster tripolium plants are rare near the sea dike but their cover and size increases further away from the dike. In the plot with 1.5 sheep/ha the height of the vegetation increases along the gradient from the dike towards the tidal flats, due to local differences in actual grazing intensity. When grazing is stopped, Puccinellia maritima is successively replaced by Festuca rubra, Halimione portulacoides and Aster tripolium. High variability of vegetation height indicates structural diversity. Patches of higher and lower vegetation correspond with the distribution pattern of different plant species. In terms of nature conservation cessation of grazing is recommended.     

Estuarine gradients and the growth and development of Agapanthia villosoviridescens, (Coleoptera), a stem-borer of the salt marsh halophyte Aster tripolium

Summary Phytophagous insects of estuarine salt marshes which live inside their host plants are not directly exposed to estuarine gradients. Host plant quality, however, may change along the estuary as a result of the direct effects of these gradients; as a consequence growth and development of endophagous insects may be influenced. The results of a study of the life cycle of Agapanthia villosoviridescens (Coleoptera, Cerambycidae), a stem-borer of the halophyte Aster tripolium, on three salt marshes along the Westerschelde estuary (the Netherlands) are in line with this hypothesis. It was shown that in upstream direction (1) mean larval weights were consistently higher during the entire period of larval development; (2) the percentage of late instars on a given sampling date generally was higher; (3) the percentage of larvae which underwent successful metamorphosis increased. Furthermore, (4) dry weight of the imagos was highest on the least saline marsh. The effects of estuarine gradients on the Aster host plants was indicated by differences in growth and chloride content between the populations of the three marshes. The non-overlapping geographic distribution of Agapanthia villosoviridescens and its host plant Aster tripolium on the Westerschelde salt marshes may be related to the effects of estuarine gradients on the suitability of the host plant.     

Drought, but not salinity, determines the apparent effectiveness of halophytes colonized by arbuscular mycorrhizal fungi

The halophytes Plantago maritima, Aster tripolium, Artemisia santonicum, Puccinellia limosa, Festuca pseudovina and Lepidium crassifolium from two different saline soils of the Hungarian steppe were examined for colonization by arbuscular mycorrhizal fungi (AMF). The salt aster (A. tripolium) and the sea plantain (P. maritima) were examined more thoroughly by recording root colonization parameters, the salt content in the soil and monthly precipitations in 2001 and 2002. Mycorrhizal colonization was maximal in late spring to early summer and had a second peak later in the autumn. Arbuscule formation and overall mycorrhizal colonization appeared to be inversely correlated with the intensity of rainfall at the investigated sites. The results suggest that, in addition to seasonality, drought may play an important role in governing mycorrhizal activity in saline habitats. In greenhouse experiments, conditions in which AMF could overcome the inhibitory effects of sodium chloride on establishing plant–mycorrhizal symbiosis were not met.     

Phytochrome et germination des semences de Rumex

The Rumex alpinus L. achenes show a special type of positive photosensitivity: several red light irradiations are necessary to induce optimal germination. The achenes deprived of their pericarp lose their photosensitivity and germinate readily in the dark. By spectrophotometry in vivo, the presence of the Pr form of phytochrome is revealed in these seeds. Short periods at extreme temperatures (30°C or 5°C) induce a good proportion of achenes to germinate. The gibberellins are inefficient on achene germination contrary to benzyladenine which exhibits some activity. A slight improvement of GA₄ effects has been detected on scarified fruits.     

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.     

Sulphide tolerance in coastal halophytes

The effect of sulphide on the growth of several species of salt-marsh plants was investigated. Relative growth rates were significantly reduced in two upper-marsh species, Festuca rubra and Atriplex patula, and in the lower-marsh species Puccinellia maritima. However the growth of Salicornia europaea, a species frequently associated with sulphide-containing sediments, was unaffected. In a separate experiment the wide ranging halophyte Aster tripolium, also appeared to be tolerant of sulphide at a concentration frequently encountered in salt marshes. Sulphide pretreatment inhibited the activity of two metallo-enzymes, polyphenol oxidase and external phosphatase, in plants from the upper marsh, but had no effect on enzymes from P. maritima or S. europaea. The rate of respiration by root tissue was significantly reduced in all of the species investigated but whereas the uptake of ⁸⁶rubidium was markedly inhibited in the other three species, uptake by S. europaea showed a significant stimulation. Similarly, whereas sulphide-grown plants of F. rubra, A. patula and P. maritima had a considerably reduced tissue iron content, the total iron concentration in S. europaea tissues was comparable to that of the controls. When the sulphide-tolerant species A. tripolium was grown in sulphide-containing media there was no significant effect on the tissue concentration of any of the elements investigated. These results are discussed in relation to possible mechanisms of sulphide toxicity and resistance.     

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     

The distribution of an intertidal aphid,Pemphigus trehernei foster,on marine saltmarshes

Summary The intertidal aphid, Pemphigus trehernei, is restricted to aster plants (Aster tripolium) growing near the edges of creeks and salt pans in low areas on tidal saltmarshes. Aphid populations in mid-marsh areas do not show an equivalent edge effect. In low-marshes the edge soil generally has more surface cracks and cavities which can accommodate aphids and a higher % air space than soil from regions away from edges. Adults and 1st instars can penetrate edge soil more readily than non-edge soil. Aphid abundance is positively correlated with % air space in low-marsh regions. Aphid populations established on potted asters were destroyed or reduced to very low levels when transferred to non-edge regions of low-marsh areas. It is suggested that aphid distribution is primarily limited by soil conditions, especially the extent of soil drainage.