Effects of temperature,salinity and cold stratification on seed germination in halophytes

Salt tolerance of halophytes corresponds with the habitat requirement of the species. It is an important factor during the germination phase and it can determine successful establishment. This paper presents the effects of alternating temperature–light regimes (4/8°C, 10/20°C, 20/32°C; 12 h dark: 12 h light) and different salinity levels (0, 200, 400, 600 mmol l²¹ NaCl) on seed germination of five halophytes, Halimione pedunculata, Bupleurum tenuissimum, Aster tripolium, Triglochin maritimum and Armeria maritima. The five species differ with respect to family and life‐form and spatially correspond to a decreasing salt gradient (i.e. distance from salt water, with H. pedunculata being the most tolerant and A. maritima being the least). Armeria maritima, A. tripolium and T. maritimum seeds were additionally subjected to a cold stratification experiment. The results showed that Halimione pedunculata, an annual therophyte of year‐round heavily saline habitats, was dormant under all experimental conditions. Bupleurum tenuissimum, a species typical to sites of varying salinity prone to leaching during spring and autumn rainfall, germinated best under cold and warm temperatures, but only under non‐saline conditions. Aster tripolium and T. maritimum, close neighbours in salt marshes, showed very similar germination behaviour: seeds of both species tolerated high levels of salinity and germinated best in summer temperatures during periods of highest soil salinity, and germination was significantly promoted by cold. Armeria maritima, a species usually found on the marginal fringes of saline habitats, germinated only under low salt levels and maximum germination was under cold (spring) and warm (autumn) temperatures, with no significant effect of cold stratification.     

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.     

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.     

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.     

Germination and early establishment of lower salt-marsh species in grazed and mown salt marsh

Abstract. Lower salt-marsh species occur more in the grazed than in the mown sites of a salt marsh in Schiermonnikoog in the Netherlands. This was demonstrated by a sowing experiment which determined which characteristic of the stand structure, height of the canopy or percentage of bare soil, is responsible for this difference. The total number of seedlings which emerged was negatively related to the height of the canopy and positively to the percentage of bare soil. The survival of seedlings of Suaeda maritima and Plantago maritima could be explained by the height of the canopy and for the latter species also by the percentage of bare soil. The survival of Aster tripolium and Plantago maritima was higher in grazed than in mown sites. Since the amount of bare soil was higher than seemed necessary for germination and survival, it was concluded that the taller canopy was responsible for the absence of lower salt-marsh species in the mown sites.     

Effect of salinity on seed germination and seedling growth of the halophyte Suaeda japonica Makino

Seed germination and seedling growth of the annual halophyte species Suaeda japonica Makino were investigated in response to variable salinity of sediment pore water. The germination percentage of S. japonica’s soft brown seeds, which are dominant among dimorphic seeds, decreased with an increase in salinity, although germination was still observed at 1200‐mM NaCl concentration. The germination percentage and germination speed observed in April were higher than those observed in December when treated with sediment water with 400–1200 mM of NaCl concentrations. These data suggest that S. japonica seedlings could be established on sediments that experience high temperatures. Germination recovery of S. japonica seeds transferred from 600‐mM NaCl containing sediment (seawater equivalent) was lowest among 0–1200‐mM NaCl treatments, implying the low tolerance of seawater conditions of S. japonica seeds. Seeds germinated in 900‐ to 1200‐mM NaCl medium showed poor growth, but survived, in hypersaline conditions, and exhibited improvement in growth upon transfer to lower salinity.     

Effect of oxygen availability and salinity on EARLY LIFE HISTORY STAGES OF SALT MARSH PLANTS. I. Different germination strategies of Spartina alterniflora and Phragmites australis (Poaceae)

Gradients in oxygen availability and salinity are among the most important environmental parameters influencing zonation in salt marsh communities. The combined effects of oxygen and salinity on the germination of two salt marsh grasses, Spartina alterniflora and Phragmites australis, were studied in growth chamber experiments. Germination of both species was initiated by emergence of the shoot and completed by root emergence. Percentage S. alterniflora germination was reduced at high salinity (40 g NaCl/L) and in decreased oxygen (5 and 2.5%). In 0% oxygen shoots emerged, but roots did not. P. australis germination was reduced at a lower salinity (25 g NaCl/L) than S. alterniflora, and inhibited at 40 g NaCl/L and in anoxia. However, a combination of hypoxia (10 and 5% O2) and moderate salinity (5 and 10 g NaCl/L) increased P. australis germination. When bare areas in the salt marsh are colonized, the different germination responses of these two species to combinations of oxygen and salt concentrations are important in establishing their initial zonation. In high salinity wetlands S. alterniflora populates the lower marsh and P. australis occupies the high marsh at the upland boundary.     

Plant community structure in reclaimed lands on the West Coast of Korea

Plant species distribution was studied on five reclaimed lands and one intertidal flat (control) on the western coast of Korea. Nineteen soil properties were analyzed. Of these, soil moisture, electrical conductivity, and levels of Na and Cl had the greatest effect on plant distribution. The plant species were divided into four groups, according to the amount of soil moisture found at their habitats.Triglochin maritimum andTypha angustata were found on the wettest sites;Phragmites communis, Carex scabrifolia, Suaeda japonica, Zoysia sinica, andSalicornia herbacea in places with relatively high moisture;Aster tripolium andPhacelurus latifolius in areas with medium levels of moisture; andAtriplex subcordata, Chenopodium virgatum, andTrifolium repens in the driest areas. The species also were divided into four groups, according to the degree of soil electrical conductivity: Highest,S. herbacea, Limonium tetragonum, Suaeda asparagoides, andS. japonica; Relatively high,A. tripolium, C. scabrifolia, P. communis, Artemisia scoparia, andZ sinka; Relatively low,Setaria viridis, C. virgatum, Sonchus brachyotus, A. subcordata, Calamagrostis epigeios, andT. angustata; and Lowest,Imperata cylindrica var.koenigii, Aeschynomene indica, Lotus corniculatus var.japonicus, andT. repens. On reclaimed land, plant species were found in zones, according to the degree of desalinization (i.e., levels of Na and Cl).     

Effects of saline‐waterlogging and dryness/moist alternations on seed germination of halophyte and xerophyte

In arid zones, precipitation distribution is extremely uneven, with saline‐waterlogging and dry–moist cycles appearing frequently, which negatively impact on seed germination and seedling establishment. The responses of two halophytes, Suaeda physophora and Haloxylon ammodendron, and a xerophyte, Haloxylon persicum, to saline‐waterlogging and dry–moist cycles were studied. The results showed that aeration increased seed germination for all species when seeds were submerged in NaCl, especially for xerophyte. Compared with S. physophora and H. ammodendron, seed germination, recovery germination, and total germination of H. persicum were much lower when seeds were submerged in 700 mm NaCl, especially for the recovery germination and total germination of nongerminated seeds when the seeds were desiccated and then transferred to distilled water. However, when the seeds were submerged in 700 mm NaCl with aeration, the seed germination, recovery germination, and total germination of nongerminated seeds transferred to distilled water increased dramatically for H. persicum. No adverse effect of desiccation was found on those values of nongerminated seeds pretreated in NaCl with or without aeration for the two halophytes. In conclusion, seeds of the two halophytes were more tolerant to waterlogging and dry–moist cycles than seeds of the xerophyte during emergence under saline conditions; these traits may be important for halophytes to survive extreme saline environments during the seed germination stage.     

Plant distribution in relation to soil properties of reclaimed lands on the West Coast of Korea

Plant species distribution was studied on five reclaimed lands and one intertidal flat (control) on the western coast of Korea. Nineteen soil properties were analyzed. Of these, soil moisture, electrical conductivity, and levels of Na and Cl had the greatest effect on plant distribution. The plant species were divided into four groups, according to the amount of soil moisture found at their habitats.Triglochin maritimum andTypha angustata were found on the wettest sites;Phragmites communis, Carex scabrifolia, Suaeda japonica, Zoysia sinica, andSalicornia herbacea in places with relatively high moisture;Aster tripolium andPhacelurus latifolius in areas with medium levels of moisture; andAtriplex subcordata, Chenopodium virgatum, andTrifolium repens in the driest areas. The species also were divided into four groups, according to the degree of soil electrical conductivity: Highest,S. herbacea, Limonium tetragonum, Suaeda asparagoides, andS. japonica; Relatively high,A. tripolium, C. scabrifolia, P. communis, Artemisia scoparia, andZ sinka; Relatively low,Setaria viridis, C. virgatum, Sonchus brachyotus, A. subcordata, Calamagrostis epigeios, andT. angustata; and Lowest,Imperata cylindrica var.koenigii, Aeschynomene indica, Lotus corniculatus var.japonicus, andT. repens. On reclaimed land, plant species were found in zones, according to the degree of desalinization (i.e., levels of Na and Cl).     

Salt regulation in halophytes

Summary Ion concentration and saturation water content were measured in various aged leaves of halophytes growing in saline soils east of lake Neusiedlersee (Austria).All species investigated showed a substantial sodium accumulation within the maturing organs accompanied by a considerable potassium decline. In most species chloride concentration rises distinctly with increasing leaf age, too, whereas concentration shifts of alkaline earth ions and of sulfate (except in Plantago maritima, Lepidium crassifolium and Crypsis aculeata) are of comparably less importance.Saturation water increases markedly in succulent species (Suaeda maritima, Chenopodium glaucum, Spergularia media, Lepidium crassifolium) and to a less degree in xerophytic monocotyledons (Puccinellia distans, Crypsis aculeata, Bolboschoenus maritimus). However, this surplus of water in older leaves is not sufficient to dilute the salt to such an extent that a rise in concentration can be prevented (except chloride in Suaeda maritima and Chenopodium glaucum).Rosette plants (Triglochin maritimum, Plantago maritima, Scorzonera parviflora, Aster tripolium) with the ability to renew their leaves continuously throughout the growth period are characterized by only insignificant changes of saturation water content with increasing leaf age. In these plants, shedding of old salt-saturated leaves is thought to be the main strategy for salt regulation.A modification of Steiner's classical concept of different salt regulation types is proposed, based on original findings about salt regulation in Austrian halophytes and on new bibliographical data upon additionally revealed regulatory principles in halophytes and saltaffected nonhalophytes.Respectfully and thankfully dedicated to Prof. Dr. H. Kinzel on the occasion of his 50th birthday.     

The effect of plant growth regulators, nitric oxide, nitrate, nitrite and light on the germination of dimorphic seeds of Suaeda salsa under saline conditions

Suaeda salsa, a leaf succulent shrub in the family Chenopodiaceae, is one of the most important halophytes in China. Suaeda salsa produces dimorphic seeds (soft brown seeds and hard black seeds). Seeds of S. salsa were collected from the coastal salt flats near Huanghua City, China. Experiments were conducted to determine the salinity-alleviating effect of plant growth regulators, nitric oxide, nitrate, nitrite and light on the germination of dimorphic seeds of S. salsa. Brown seeds had a higher germination rate than black seeds in all experiments. Black seeds were more sensitive to salt in the absence of light in comparison to brown seeds. Brown seeds absorbed water more quickly in comparison to black seeds and were found to be more tolerant of salt stress. Our results showed that 1-aminocyclopropane-1-carboxylate (ACC, the immediate precursor of ethylene), nitrite, GA₄ and BA improved seed germination in the presence of salt. However, nitrate, GA₁, GA₃ failed to alleviate salt stress. ABA inhibited seed germination and seedling growth. Possible mechanisms involved in the alleviation of salt stress in S. salsa seeds and the ecological adaptation of the seeds to the environment are discussed.     

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.     

The sediment burial depth and salinity control the early developments of Suaeda salsa in the Yellow River Delta

Sediment deposition is a common phenomenon in the estuary area. Pot control experiments were conducted to evaluate the interaction effects of sediment burial depth and salt stress on the seed germination and early seedling growth of Suaeda salsa (L.) Pall., an pioneer species of tidal wetland near the Yellow River Delta. The results showed that the percentage of seedling emergence, seedling emergence rate, seedling height, branch number, shoot biomass and root biomass were all significantly affected by salt stress and sediment burial depth. While the interaction of salt and burial depth significantly influenced the branch number, leaf biomass, shoot biomass and total plant biomass. Only 5 cm burial depth without salt stress should 6.25 ± 3.61% seedlings emergence. With the increasing of sediment burial depth and salt stress, percentage of seedling emergence, seedling emergence rate and plant height decreased significantly. However, under the salt treatment of 0 and 1%, the branch number increased dramatically with the increasing of sediment burial depth from 0 to 3 cm. The ratio of leaf to total biomass increased with increasing of burial depth, on the contrary, the ratio of root to total biomass decreased. 0–1 cm sediment burial depth was proved the suitable depths for seed germination of S. salsa in the coastal wetland of the Yellow River Delta. Our findings contribute to a better understanding of how to improve the seedling establishment of S. salsa under the dynamic changes of sediment deposition and salinity in the coastal wetland of the Yellow River Delta.     

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.     

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.     

Seed germination of the halophyteSuaeda japonica under salt stress

Suaeda japonica Makino belonging to the family Chenopodiaceae, is a halophyte and grows at the shore of Ariake sea in Japan. This plant presumably possesses high salt resistant nature, thus, we examined the mechanisms of seed germination under salt stress. The seeds maintained 80% germination rates on the medium containing 0.7 M NaCl. Germination rates varied depending on salt type; the germination rates under NaCl or KCI exhibited relatively lower values than ones under sodium gluconate or potassium gluconate. This different responses for salts seemed to be as a result of the presence of Cl⁻ ions. Although very high levels of betaine (compatible solute), were kept in the seedlings grown under no salt stress, the contents gradually increased as concentration of NaCl increased. Betaine is a factor present in plants that works to alleviate the effects of excessive soil salts. It is synthesized in leaves from betaine aldehyde, and this process is catabolized by betaine aldehyde dehydrogenase (BADH). When the seedlings were cultivated on the medium without NaCl, relatively high level of BADH activity was found. The activity increased 5-fold in the seedlings grown under 0.5 M NaCl stress. Increases in betaine content and BADH activity were found during seed germination. InS. japonica, the salt stress promoted BADH activity, subsequently endogenous betaine contents were increased, and increased betaine seemed to secure seed germination under salt stress.     

Plant succession and interaction between soil and plants after land reclamation on the west coast of Korea

Plant succession and the interaction between soil and plants after reclamation were investigated on the west coast of Korea. Our study included one natural tidal flat site (Namdong, 3 km²), and five sites that differed in the number of years since being reclaimed: Hyundai A, 6 km² (1 yr); Hyundai B, 5 km² (2 yr); Jangdeog, 5 km² (8 yr); Mado, 2.5 km² (12 yr); and Baegseog, 1 km² (30 yr). The number of plant species occurring at each site was 10, 9, 15, 30, 28, and 38, respectively. Based on distribution ranges of the plants along gradients of salt and moisture contents in the soil, and species associations and life forms, plant succession was divided into two sere groups, hydrarch and xerarch. The major species of the former werePhragmites communis andTypha angustata, but the sequence of the sere could not be identified. Species associated with the latter were [Suaeda japonica] → [Salicornia herbacea, S. japonica, Atriplex subcordata, Suaeda asparagoides] → [Aster tripolium, Carex scabrifolia, Zoysia sinica, Limonium tetragonum] → [Artemisia scoparia, Calamagrostis epigeios, andSetaria viridis] → [Imperata cylindrica var.koenigii, Sonchus brachyotus, S. viridis] → [Aeschynomene indica, Lotus corniculatus var.japonicus, Trifolium repens, and other non-halo-phytes]. In certain circumstances, the first and second stages replacedS. japonica withC. scabrifolia andZ sinica. The progression of interactions between soil and plants through succession was salt leaching → increase in species richness and biomass → increase in soil organic matter → increase in total nitrogen and decrease in bulk density of soil, and/or salt leaching → increase in phytomass → decrease in soil-available phosphorus.     

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.