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This master project has a focus on extraction, separation and identification of flavonoids and other phenolic compounds in Plantago maritima. Previously in Norway there has only been done one small pilot study (bachelor project in Jordheim group) on the phenolic compounds in this plant using only DAD-HPLC, so the Norwegian polyphenol profile of this species isn’t well documented. The polyphenolic profile of a species can show difference within geographical gradients due to different external parameters such as temperature, drought etc., normally this is seen for the quantitative amounts found. Qualitative polyphenolic profile variations are observed, however not to the same extent. Based on a literature survey, almost forty different polyphenols/phenolics are found in the genus in the classes of phenolic acids (cinnamic acid and hydroxybenzoic, and related esters), phenylethanoids and flavonoids. Both unsubstituted aglycones, mono and di-glycosylated compounds are seen. From literature ten polyphenols have been found in P. maritima: Caffeic acid (a), p-coumaric acid (b), ferulic acid (c), luteolin (14), chrysoeriol, diosmetin (17), apigenin-7-O-glucoside (2), luteolin-7-O-glucoside (3), verbascoside (10) and plantamajoside (1). In literature these ten compounds are confirmed through DAD-HPLC, ToF-MS (a, b, c, 14, 1 and 10) and MS/MS (2, 3 and 14). Plantago maritima samples were harvested in Tømmervågen (60.440802°N, 5.302253°E) Bergen 23.06.2019 and initial DAD-HPLC analysis was performed. Extraction was performed with aqueous methanol (60:40), and extracts were combined and evaporated. The crude extract was purified with XAD-7, and finally with preparative HPLC. The different stages of purity and fractions were analyzed with DAD-HPLC and LC-MS in negative ESI mode. Selected preparative HPLC fractions were run on NMR. In addition, crude and purified fractions have been extensively analyzed with DAD-HPLC (UV-Vis and retention time) and LR-LC-MS (including MS/MS on glycosylated flavonoids). A total of 18 phenolics were identified in Norwegian P. maritima. Only trace amounts were found from caffeic acid (a), p-coumaric acid (b) and ferulic acid (c). Plantamajoside (1) and luteolin (14) were confirmed with NMR (1D, 2D) analysis. These have previously only been tentatively identified in the species using HPLC, ToF-MS (1 and 14) and MS/MS (14). The flavonoid tricin (18), previously found only in P. phaeostoma, was identified with co - chromatography on HPLC and LC-MS using a tricin standard. A tricin glucoside (11) has also been identified (by LR-LC-MS and MS/MS) earlier found in P. crassifolia. Apigenin glucoside (2), luteolin 7 – O – glucoside (3), verbascoside (10) and diosmetin/chrysoeriol (17) were confirmed to be present (HPLC, LC-MS and MS/MS on 3 and 11). Ten compounds were for the first time in addition tentatively identified to be present within P. maritima: Campneoside I (4), nepitrin (6 – methoxy luteolin 7- glucoside) (6), luteolin glucuronide (7), quercetin arabinoside (8), tricin glucoside (11), rosmarinic acid (12), Apiin (apigenin 7 – apioglucoside) (13), martynoside (15), apigenin (16) tricin (18). (HPLC, LC-MS and MS/MS on 6, 7, 11 and 13). These have been found in the genus before by using DAD-HPLC, NMR (1, 3, 10, 11 and 15), TLC (12), MS/MS (1, 2, 4, 10, 14, 15, 16 and c), HR-MS (1, 2, 4, 10, 14, 15, 16 and c) In addition, two components tricin diglucoside (5) and diosmetin rutinoside (9) (HPLC, LC-MS and MS/MS on 5) were tentatively identified in both the species and in the genus for the first time. Masteroppgave i kjemi KJEM399 MAMN-KJEM

Several methods of tillage are among the practices most destructive to inland saline grasslands, and can result in their complete deterioration. Despite the high frequency of habitat destruction by these means, the recovery of the saline vegetation and regeneration abilities of halophytes are still not sufficiently known. The reaction of inland saline grasslands to disk tillage was studied on the Jatov site located in the Pannonian bioregion of Central Europe. Changes in species composition were tested using ordination methods (DCA, CCA). Although the species composition recovered to a certain extent after the 4 years following the disturbance, the original zonation of halophytic vegetation was disturbed and did not recover. Disturbance primarily suppressed perennial halophytes and promoted the occurrence of species with a ruderal strategy. Testing of halophyte reaction to tillage showed the contracting responses of various halophytic species. The species Atriplex littoralis, Plantago maritima and Matricaria recutita reacted positively to tillage due to the reduced competition. The species Artemisia santonicum subsp. patens and Dichodon viscidum were suppressed by tillage in initial stages, but later recovered their populations. Only A. santonicum subsp. patens was able to return to the abundance seen before the disturbance. The only halophytic species which was unable to recover was a shallow-rooted grass Puccinellia distans. Our study showed that a Pannonian alkali steppe is able to recover quite quickly after tillage, if the soil disturbance is shallow and subsequent management takes place. Some halophytes may even profit for a short time from tillage, but the original species composition cannot be restored in the short term.

The presented study supplements the knowledge on ion-exchange capacity, swelling capacity (elasticity) of the plant cell wall, and the accumulation of heavy metals in halophytic species Plantago maritima and Triglochin maritima in the tidal zone of the White Sea western coast. The littoral soils of the coastal territories are sandy or rocky-sandy, medium and slightly saline with poor content of organic substances, Mn, Zn, Ni, and Pb. Studied soils are considered as uncontaminated by heavy metals because they contain background amounts of Fe and Cu. Sea water is significantly polluted by Fe (3.8 MPC) and Ni (55 MPC), has poor content of Zn and Cu and background level of Pb and Mn. The coastal dominant plant species P. maritima and T. maritima were characterized by intensive metals accumulation which was reflected in the coefficient of biological absorption (CBA) of metal by a whole plant. For P. maritima the following metal accumulation series was obtained: Cu (3.29)> Zn (2.81)> Ni (1.57)> Pb (1.30)> Mn (1.21)> Fe (0.97), and for T. maritima: Ni (3.80)> Fe (2.08)> Cu (1.91)> Zn (1.84)> Pb (1.51)> Mn (1.31). Roots accumulated 50–70% of Ni, Cu, Zn, Pb and Mn of the total metal content in the plant while leaves and stems contained 30–50%. Fe was allocated mainly in the roots (80%). The ion-exchange capacity of the plant cell wall for P. maritima and T. maritima was established as follows correspondingly: 3570–3700 and 2710–3070 μmol g-1 dry cell weight per leaf; 2310–2350 and 1160–1250 μmol g-1 dry cell weight per root.

Structural and functional traits in leaves of sea plantain (Plantago maritima L.) inhabiting intertidal and supratidal zones of the White Sea coast were studied. At the intertidal, the plants were found to have smaller and thicker leaves with more frequent stomata and thinner cover tissues in comparison with the plants collected from the supratidal. Leaf fluorescence and stomatal conductance obeyed the circadian rhythm in plants residing in the supratidal. The tested parameters considerably changed according to tidal dynamics in the intertidal plants. Two stable states (high and low tides) and two transition states (while water level ascends or descends) were defined. In both stable states, the plants demonstrated high functional parameters of fluorescence (Fv/Fm, Y(II), and ETR) and open stomata. In the transitional states, the stomata were partly closed, and the functional activity was inhibited. Leaf structural peculiarities, leaf gas films, the inducible CO2-concentrating mechanism, and the determining role of the stomata are considered in terms of the hypotheses explaining the responses of sea plantain to tidal dynamics on the White Sea coast.

QUESTION: Studies in the northwest Atlantic have shown that nutrient enrichment can severely impact salt marshes, undermining their broad range of high‐value ecosystem services. However, biogeographical differences in plant communities may preclude extrapolation of these findings to other regions. The few investigations of such impacts elsewhere, including in Europe, have been strongly localised, restricting generalisations, and have not addressed community‐level properties. LOCATION: South and east coasts of Ireland (northeast Atlantic). METHODS: We conducted a field survey of vascular plants and soil properties in fifteen salt marshes representing broad environmental gradients but comparable biogeographical settings, to test if salt marsh plant (a) diversity and (b) community composition are related to soil nutrients. RESULTS: Species richness and Shannon diversity were both negatively related to soil NOₓ⁻, and Shannon diversity had a hump‐shaped relationship to labile P. Multivariate community composition was highly significantly related to NOₓ⁻, to a lesser degree to P, and marginally significantly to NH₄⁺. This was most clearly evident in the positive association of the ground cover of an evergreen dwarf shrub, Atriplex portulacoides, and a weaker negative association of a forb, Plantago maritima, with NOₓ⁻, indicating competition between these key and functionally contrasting mid‐salt‐marsh species. The relationship between community composition and P was most evident through the positive association of P with the invasive cordgrass, Spartina anglica. CONCLUSIONS: Physiognomic and functional differences between the herb‐rich Pl. maritima community and monospecific A. portulacoides stands likely have implications for ecosystem services, as would a continued expansion of S. anglica. Further studies to ascertain causality could determine whether such transitions can be driven by nutrient enrichment.

International audience; This article presents a case study of recolonization by flora constituting a continental salt meadow type vegetation after the removal of an embankment on a continental salt marsh in the Limagne plain (Puy-de-Dôme, France). Salt marshes indicator plants was monitored in presence/absence in a network of 104 meshes of 10 m². We were able to show the rapid and lasting colonization of Puc-cinellia distans and Juncus gerardi, but also the slower and variable installation of Spergula media and Plantago maritima. We mapped spatial and temporal evolutions of this colonization for 6 different species. Among the halotolerant species, one should note the increase in the population of endan-gered species such as Inula britannica. The monitoring protocol allows us to conclude on the interest of this type of embankment removal operation to restore the abiotic parameters and the flora of the «continental salt meadow» vegetation.; Cet article présente une étude de cas de recolonisation par la végétation de type pré salé continental après des travaux de suppression d’un remblai sur un marais salé continental dans la plaine de la Limagne (Puy-de-Dôme, France). Les plantes indicatrices des prés salés ont été suivies en présence/absence dans un réseau de 104 mailles de 10 m². Nous avons pu montrer la colonisation rapide et durable de Puccinellia distans et Juncus gerardi, mais aussi l’installation plus lente et variable de Spergula media et de Plantago maritima. Nous avons cartographié les évolutions spatiales et temporelles de cette colonisation pour 6 espèces différentes. Parmi les espèces halotolérantes, on notera l’augmentation de la population d’espèces menacées comme celle d’Inula britannica. Les résultats permettent de conclure sur l’intérêt de ce type d’opération de suppression de remblais pour restaurer les paramètres abiotiques et de la capacité de la flore de se réinstaller même après vingt-cinq ans pour reconstituer une communauté végétale caractéristique des « prés salés continentaux ».

The growth conditions of halophytic species Triglochin maritima and Plantago maritima on tidal zone of western coast of the White Sea were studied. The littoral soils were characterized by medium and slightly saline, poor content of organic substances, manganese, zinc, nickel, lead, and content of natural amounts of iron and copper. Sea water was contaminated with iron (3.8 of MPC) and nickel (55 of MPC), and contained poor of zinc and copper as well as natural level of lead and manganese. The studied plant species actively accumulated metals. For T. maritima the following metal accumulation series was obtained: Ni (3.80) > Fe (2.08) > Cu (1.91) > Zn (1.84) > Pb (1.51) > Mn (1.31) and for P. maritima, Cu (3.29) > Zn (2.81) > Ni (1.57) > Pb (1.30) > Mn (1.21) > Fe (0.97). The roots accumulated 50%–70% of Ni, Cu, Zn, Pb, and Mn. The results show that the test plants can be used in the phytoremediation process of marine territories.

Pētījuma mērķis bija veģetācijas eksperimentā kontrolētos apstākļos salīdzināt divu jūras piekrastei specifisku ceļteku sugu (Plantago coronopus un jūrmalas ceļteka) izturību pret pieaugošām NaCl koncentrācijām un jonu uzkrāšanās īpatnības. Augus audzēja automatizētās siltumnīcas apstākļos veģetācijas traukos. Abu ceļteku sugu augus apstrādāja ar pieaugošas Na+ koncentrācijas devu, katrā variantā bija pieci atkārtojumi. Eksperimenta beigās augu sadalīja morfoloģiski atsevišķās daļās, noteica to svaigo un sauso masu, ūdens satura, kā arī mērija Na+ un K+ jonu koncentrāciju, elektrovadītspēju un osmotisko aktivitāti. Abas ceļteku sugas spēja augt un vairoties jūras ūdens sāļuma apstākļos. Plantago coronopus ir uzskatāma par sāls tolerantāku sugu, nekā jūrmalas ceļteka. Abas ceļtekas ir K+ izslēdzošas un Na + jonu uzkrājošas sugas, kur galvenokārt Na+ joni tiek izmantoti osmotiskajā regulācijā kā osmotiski aktīvs aģents., The aim of the study was to compare two coastal specific plantain species (buckshorn plantain and sea plantain) to increasing NaCl concentrations and ion accumulation characteristics under controlled conditions in a vegetation experiment. Plants were grown in automated greenhouses conditions in vegetation pots. Plants of both plantain species were treated with increasing doses of Na+ concentration, there were five repetitions in each variant. At the end of the experiment plants were divided into morphologically separate parts, fresh and dry mass was determined as well water contents was measured, Na+ and K+ ion concentration, electrical conductivity and osmotic activity were also tested. Both plantain species were able to grow and reproduce under seawater salinity. Buckshorn plantain was considered to be a more salt-tolerant species than sea plantain. Both plantain species are K+ excluding and Na+ ion accumulating species, where mainly Na+ ions are used in osmotic regulation as an osmotically active agent.

The near elimination of inland salt marshes in Central Europe occurred throughout the 19th and 20th centuries, and the currently remaining marshes exist in a degraded condition. This work examines the impact of groundwater level on the growth of plants from a seed bank obtained from a degraded salt marsh in proximity to still existing one through an ex-situ experiment. An experimental tank was set up with the sample seed bank experiencing differing levels of water level. There were 1233 specimens of 44 taxa grown from the seed bank, of which 5 species were abundant, and 10 species are considered as halophytes. Only Lotus tenuis from halophytes was more abundant, and only five species of halophytes were represented by more than three individuals. The water level has a significant impact on the number of species (based on linear regression analysis) as well as species distribution among different water level treatments (a non-metric multidimensional analysis (nMDS) followed by linear regression). The results show a strong negative relationship between the average water level and the number of species. The water level did not affect the species composition of halophytes, but differences in individual species abundances were found among the halophytes. The species Bupleurum tenuissimum, Crypsis schoenoides, Melilotus dentatus, and Plantago maritima grew on the drier and non-inundated soils. Tripolium pannonicum, Spergularia maritima, and Lotus tenuis grew on both wet and dry soils. Trifolium fragiferum and Bolboschoenus maritimus were found in places with water stagnant at the soil level. Pulicaria dysenterica grew in inundated soil.

We examined whether the presence or absence of different environmental stressors influenced the reproductive potential of a saltmarsh species—Plantago maritima. We focused on total seed output, seed quality, and biomass of progeny. So far, there are no studies trying to answer the question of how different saltmarsh management affects the quality of seed in saltmarsh species. For the purposes of the study, plots subjected to light mowing, light or heavy grazing, trampling, or rooting were designated in three nature reserves in Poland. On each plot, the abundance of infructescences per sq. meter was calculated. Mature infructescences were collected, and their length and number of fruit capsules were measured. The seeds obtained from fruit capsules were weighted and sown in controlled conditions. The germination rate and the final germination percentage were calculated. A representative number of sprouts were grown. After a period of 2 months, the individuals of P. maritima were harvested and their total dry mass was measured. It was found that heavy grazing had the greatest effect on all of the studied characteristics. The presence of this factor resulted in shorter infructescences with a smaller number of fruit capsules. However, this phenomenon was compensated by the higher abundance of infructescences per sq. meter. At the same time, seeds produced by grazed individuals were significantly lighter. Interestingly, intensive trampling by people affected P. maritima individuals in a similar way to heavy grazing, while mowing and rooting had less impact on the considered characteristics. Although a positive correlation between seed mass and germination success was found, the altogether lower seed mass had a negligible effect on germination parameters. Also, the differences in seed parameters did not affect dry mass of obtained progeny grown in laboratory conditions. Synthesis and applications: Different environmental stressors, such as grazing and mowing, have an effect on reproductive potential of a saltmarsh species P. maritima. In the case of habitats created anthropogenically, such as brackish saltmarshes, the role of management is crucial for their conservation. Therefore, searching for the best active protection methods is important. In light of the results obtained, extensive or rotational grazing appears to be the best form of saltmarsh management., In the case of habitats created anthropogenically, such as brackish saltmarshes, the role of management is crucial for their conservation. Therefore, searching for the best active protection methods is important. In light of the results obtained, extensive or rotational grazing appears to be the best form of saltmarsh management

Nowadays, it is relevant to study the mechanisms of rhythmic processes that are the basis of the vital activity of all organisms. From this point of view, it is necessary to study the functioning of organisms living in tidal conditions on the coastal zone of the northern seas. Twice a day, plants are flooded with water on the intertidal zone, and then they go out for drying—there is a change in the water and air environment, which is accompanied by changes in light, temperature, oxygen and CO2 concentrations, pressure, and other parameters. These conditions have led to the formation of a unique halophyte complex of higher terrestrial plants, which have formed various adaptive mechanisms that provide the possibilities for normal existence. Three zones are distinguished on the coastal zone of the White Sea: supratidal, intertidal, and subtidal zones. For each zone, the following species was investigated: for supratidal—Alopecurus arundinaceus, Atriplex glabriuscula, Carex subspathacea; for intertidal—Tripolium vulgare, Triglochin maritima, Salicornia europaea, and three species of Plantago genus; for subtidal—Zostera marina. Species growing on intertidal and subtidal zones are resistant to flooding and have various structural and functional adaptations. The photosynthetic characteristics (CO2 rate of gas exchange, transpiration, stomatal conductivity, and CO2 concentration in the intercellular spaces of the leaves) were studied using the LCPro + portable gas analyzer at natural light intensity and air temperature. Analysis of CO2 exchange curves by the model of Farquhar et al. showed a difference in the values of the maximum carboxylation rate (Vcmax), the electron transport rate at light saturation (Jmax), the utilization rate of triose phosphate, and a number of other parameters in plants of different zones. A comparative study of physiological parameters showed the highest functional activity in plants in the intertidal zone, which undergo daily tidal dynamics. The influence of environmental factors on the content of chlorophylls a and b and the amount of carotenoids was estimated. Evaluation of the implementation of photosynthetic activity in natural habitats in relation to the maximum values at saturating concentration of CO2 showed that the species of higher vascular plants realize their potential in the range of 6–20% on the supratidal zone, from 10 to 70% on the intertidal zone, and from 30 to 60% on the subtidal zone. In the case of a global increase of CO2 concentration, we can expect a significant increase in CO2 absorption by the following halophytes: T. vulgare, Plantago maritima, Z. marina, Al. arundinaceus, and At. glabriuscula, as well as an increase in productivity of P. maritima and Z. marina among the studied halophytes of the coastal zones of the northern Holarctic seas.

1.Halophyte distributions on saltmarshes are strongly related to elevation in the tidal frame. However, collinearity between elevation, the consequent inundation regime, and sediment waterlogging/redox potential obscures the proximate causes of distribution patterns. We sought to distinguish the effects of elevation per se from those of waterlogging by manipulating microtopography. 2.We experimentally manipulated elevation by ±15 cm at locations that spanned the elevation ranges of three saltmarshes recently reactivated by managed coastal realignment. Experimental plots were initially cleared of any vegetation. Elevation and sediment redox potential were determined for each plot. We planted five perennial species (Armeria maritima, Atriplex portulacoides, Limonium vulgare, Plantago maritima and Triglochin maritima) in half of the plots, recording survival over four years, and monitored natural colonisation of the other plots. 3.Overall, redox potential increased with elevation. Sediments were more oxidising in raised plots and more reducing in lowered plots. Redox reductions in lowered plots were in line with those that would be predicted from the overall redox/elevation relationship, but increases in raised plots were greater than predicted from elevation alone. 4.Plant colonisation and survival was poorer in lowered plots and, for most species, improved in raised plots. This poorer colonisation and survival can, in part, be attributed to the concomitant alterations in redox potential and elevation in the tidal frame, but microtopographic manipulation also had substantial independent effects on plant performance, including on the survival of all planted species and the colonisation of Puccinellia maritima, Salicornia europaea agg. and Tripolium pannonicum. 5.Synthesis: Microtopography can have effects on sediment chemistry and plant performance similar in magnitude to those of overall tidal elevation. Understanding how its effects modulate the relationship between tidal elevation, redox and other environmental conditions helps clarify the abiotic factors that fundamentally determine halophyte colonisation and survival. These results support the use of topographic manipulation to enhance the diversity of created saltmarshes.

The multifunctionality and delivery of ecosystem services from green roofs is improved by biological diversity of the roof vegetation. However, the frequency and intensity of drought episodes on extensive green roofs may limit the use of non-succulent species and the potential functional and phylogenetic diversity of the vegetation. Wind accelerates water use by plants and desiccation of the green roof substrate, and may be a key factor in selection of non-succulent plant species for green roofs. In this study, we tested wind interactions with green roof substrate composition and the effects on plant and substrate water balance, overall plant performance, and wilting and survival of three non-succulent species (Plantago maritima L., Hieracium pilosella L., and Festuca rubra L.) under realistic prolonged water deficit conditions. We found that, regardless of species or substrate tested, wind accelerated drought response. Drought-stressed plants exposed to wind wilted and died earlier, mostly due to more rapid desiccation of the growth substrate (critical substrate moisture content was 6-8%). The moderate wind levels applied did not affect plant performance when not combined with drought. Species with contrasting growth forms showed similar responses to treatments, but there were some species-specific responses. This highlights the importance of including wind to increase realism when evaluating drought exposure in non-succulent green roof vegetation.

The paper presents the results of geobotanical studies on the distribution and resources of Aster tripolium L. and Plantago maritima L, two rare halophytes in Poland. The research was conducted in northern Poland, along the Baltic coast in 2013. The present distribution of the two species was compared with historical data and general trends of and threats to these two species were examined. In total, 33 sites of A. tripolium and 18 of P. maritima were found in the research area. The resources of both species have been perceptibly depleting during last 150 years, which is mostly due to human agencies (e.g. habitat devastation caused by growing urban areas and the change in management and/or habitat condition). In order to preserve both species, it may be necessary to start an ex situ conservation program.

Seven species not previously reported for Lebanon and one species re-located after 183 years are reported here after their discovery during an ex situ seed conservation project in the country (Bombycilaena erecta (L.) Smoljan., Centaurea diffusa Lam.; Hypochaeris radicata L.; Gypsophila perfoliata L.; Euphorbia cheiradenia Boiss. & Hohen.; Stachys setifera C.A.Mey.; Stachys zoharyana Eig and Plantago maritima L.).

The phytosociology and ecology of Scorzonera parviflora stands were studied in the Pannonian Basin and Western Balkans. In total, we used 60 relevés (46 collected for this study and 14 from the literature) for the ordination analysis. Optimal conditions for S. parviflora are species-poor coastal habitats in regions with hot summers and with high soil salinity and nutrient-richness. The species reached high cover values in most cases (25–50%) and it often dominated the stand. The classification of phytosociological data shows four floristically and ecologicaly well characterized clusters. Cluster 1 was identified as the rare association Junco maritimi-Scorzoneretum parviflorae, sampled only in coastal Croatia (on the island of Pag) and inland Hungary (Fertőszéplak). Cluster 2 groups meadow-like salt marsh vegetation in transition between two vegetation types, Limonio narbonensis-Puccinellietum festuciformis and Trifolio-Hordeetum secalini. Cluster 3 represents the typical variant of the Scorzonero parviflorae-Juncetum gerardii, while cluster 4 was determined as a variant with Tripolium pannonicum of the Scorzonero parviflorae-Juncetum gerardii characterized by a higher presence of obligate halophytes (Tripolium pannonicum, Puccinellia distans agg. and Plantago maritima) than in cluster 3. Our study shows that vegetation with Scorzonera parviflora in inland Central and SE Europe belongs to the Juncion gerardii and stands of coastal Croatia are related to the Juncion maritimi alliance. The species composition and the classification of the examined stands in the Pannonian and Transylvanian basins were influenced more by abiotic environmental factors than by current management.

Our study is focused on native spontaneous species of saline ecosystems Plantago maritima . Plants were cultivated at several salt concentrations (0, 50, 100, 200, 300, 400 and 500 mM NaCl) in a glass greenhouse under semi-controlled conditions. Growth parameters, water parameters and ionic status were determined and they were used as criteria to assess the response of P. maritima under a salinity gradient. Catalase, guaiacaol and ascobate peroxidase activities, total protein and proline were also determined. Our results show that P. maritima is a facultative halophyte capable of expressing its maximum growth potential at relatively low concentrations of salt (less than 3 g l −1 NaCl). At high doses of salt (concentrations > 200 mM), the decrease in the growth of P. maritima is associated to a decrease in the uptake of K + . There is a disruption of the water intake of their organs and therefore results an invasion of the cytoplasm by Na + toxic ion. However, stressed plants use K + more sparingly. They invest especially in the production of biomass expressed by the dry weight of the shoots, and they use Na + and proline for osmotic adjustment. The halophyte studied is able to accumulate high levels of proline in response to increasing salt concentration. The accumulation of the amino compound, mainly in roots, is interpreted as an indicator of salt tolerance. Additionally, a significant correlation between the tolerance of the plants to salinity and the activity of several antioxidant enzymes has been observed. Hence, we suggest the possibility of using these activities as a biochemical indicator for salt tolerance in P. maritima . Our study points out two types of biomarkers of salt exposure: enzymatic biomarkers in the leaves and proline content in the roots. Both did show very good correlation with salt exposure, and thus may be considered good biomarkers of exposure with a very good dose–response relationship.

Darba mērķis bija veikt daudzfaktoru eksperimentus un aprobēt veģetācijas pētījuma metodiku jūras piekrastes reto un aizsargājamo augu sugu fizioloģiskajiem pētījumiem, lai noskaidrotu dabisko faktoru iespējamo ietekmi. Piekrastes zonas ir dinamiski mainīgas vietas, kurās sastopamas daudzas endēmas augu sugas, no kurām liela daļa ir retas un aizsargājamas. Eksperimentos izmantoja četras jūras piekrastes retās un aizsargājamās augu sugas - smiltāja neļķi (Dianthus arenarius), jūrmalas pērkonamoliņu (Anthyllis maritima), jūrmalas ceļteku (Plantago maritima) un jūrmalas āžloku (Triglochin maritimum). Augu sugas pavairoja ar sēklām vai veģetatīvi un atkarībā no sugas ekoloģijas, veica piekrastes teritorijām raksturīgas ietekmes faktoru apstrādi – apbēršanu ar smiltīm, nobradāšanu, slāpekli fiksējošu gumiņbaktēriju pievienošanu, augsnes minerālelementu un sāļu sastāva izmaiņas, vai dažādas iepriekšminēto apstrāžu kombinācijas. Visiem augiem veica hlorofila a fluorescences mērījumus, hlorofila koncentrācijas mērījumus, jonu kvantitatīvu noteikšanu un galējo morfoloģisko datu iegūšanu. Eksperimentā ar A. maritima papildus noteica arī elektrolītu noplūdes, gvajakola peroksidāzes un polifenoloksidāzes aktivitātes analīzi. D. arenarius spēja atjaunoties pēc ilgstošas apbēršanas un nobradāšanas intensitātes. Viegla apbēršana (1.3 cm smilšu slānis) veicina virszemes daļu masas pieaugumu. A. maritima augu augšanas un atjaunošanās spēju ietekmēja slāpekli fiksējošo gumiņbaktēriju klātbūtne un smilšu apbēršanas intensitāte (2 cm un 4 cm smilšu slānis). P. maritima spēja ilgstoši paciest 1000 mM sāļumu, saglabājot aktīvu fotosintēzi, bet 2000 mM augi pakāpeniski gāja bojā. T. maritimum neuzrādīja toksiskuma pazīmes dažādu jonu izmantošanai pat augstās koncentrācijās. Eksperimentus veica no 2016. gada augusta līdz 2017. gada aprīlim Latvijas Universitātes Bioloģijas fakultātes Augu fizioloģijas katedrā, LU Dabaszinātņu akadēmiskā centra siltumnīcā. Eksperimentus veica VPP EVIDEnT, „Latvijas ekosistēmu vērtība un tās dinamika klimata ietekmē’’., The aim of the study was to perform multifactorial experiments and to approve a method of vegetation experiments for investigation of impact of natural environmental factors on rare and endangered coastal plant species. Sea coast is a dynamically variable habitat, where high number of endemic plant species can be found, many of them being rare and endangered. Four rare and endangered coastal plant species were used in the present study: Dianthus arenarius, Anthyllis maritima, Plantago maritima and Triglochin maritimum. The plant species were propagated by seed or vegetatively and, according to their ecology, the main environmental factors on a sea coast were used as treatments: sand burial, trampling, application of nitrogen fixing root-nodule-forming bacyeria, mineral nutrient disbalance, and salt changes in the soil, or combinations of the previously mentioned treatments. Chlorophyll a fluorescence, chlorophyll concentration, ion quantitative measurments and final morphological data were collected from all species. In the experiment with A. maritima electrolyte leakage, guaiacol peroxidase activity and polyphenol oxidase enzyme activity was measured as well. D. arenarius recovered after prolonged period of time of sand burial and trampling. Light sand burial (1.3 cm sand layer) increased weight of the above ground parts. Nitrogen fixing root nodules and intensity of the sand burial (2 cm and 4 cm sand layer) affected growth and recovery of A. maritima. P. maritima withstanded 1000 mM salinity for a prolonged period of time, maintaining active photosynthesis, but it gradually perished at 2000 mM salinity. T. maritimum did not show any toxicity symptoms at different high ion concentrations. The experiments were carried out from August 2016 till April 2017 at the Faculty of Biology, the Department of Plant Physiology, University of Latvia in the greenhouse of the Academic Centre of Natural Sciences. The study was carried out within the project from NRP EVIDEnT „The value and dynamic of Latvia’s ecosystems under changing climate”.

We studied the vegetation development in the Pannonian salt steppes in SW Slovakia (class Festuco-Puccinellietea) for eight years on permanent plots. The saline grassland was ploughed in 2002 to cultivate wheat and barley, and after two years it was left fallow. An open vegetation structure was typical for the initial stages, dominated by the obligate halophyte Tripolium pannonicum, accompanied by Atriplex littoralis and other annuals. Species richness significantly increased in the subsequent years. From the fifth year of the study, a significant increase of facultative halophytes (e.g. Lotus tenuis, Tetragonolobus maritimus, Agrostis stolonifera) and colonization of some expansive species (Phragmites communis, Calamagrostis epigejos) was observed and accompanied by a decrease in the cover of obligate halophytes (T. pannonicum and Puccinellia distans). Plantago maritima exhibited an opposite trend, and it was the only halophyte that was able to increase its population after the disturbance, due to vegeta...

The chemical form and content of available nitrogen (N) in salt marsh substrates varies considerably. On the western coast of Ireland, habitats designated as Ombrogenic Atlantic salt marshes were formed on ombrogenic peat substrate. The peat substrate in these systems has three times more ammonium than substrate from adjacent salt marsh habitats on sand and mud substrate. This study examined the extent to which the high concentration of ammonium in peat salt marsh substrate influences the N- assimilating enzyme activity of halophytes and the extent to which N metabolism differs between species. Specifically, this work investigated whether plants from peat salt marshes are more likely to assimilate ammonium than plants from non-peat substrates. Four halophyte plant species—Armeria maritima, Aster tripolium, Plantago maritime, and Triglochin maritime—were sampled from various saltmarsh habitats including three sites on peat substrate and three on non-peat substrate, comprising sand, mud and sand/mud. The activities of N-metabolising enzymes—glutamine synthetase (GS), glutamate synthase, glutamate dehydrogenase (GDH), and nitrate reductase (NR)—were quantified in shoot and root parts. Root GS activity in Armeria maritima and shoot GS activity in Triglochin maritima were positively correlated with increasing soil ammonium levels. Root NR activity in Aster tripolium and shoot NR activity in Plantago maritima were significantly higher in plants grown on non-peat substrates than peat substrates. The shoot : root GS activity ratio in Triglochin maritima on peat substrate was more than double the ratio on non-peat substrates. It is concluded that all species tested displayed differences in N-metabolising activities depending on the chemical form and/or concentration of N in the substrate, while three out of the four species were capable of taking advantage of the high levels of ammonium in peat substrates.

Conceptual models and some empirical studies in plant systems show that species interactions can shift from competitive under low stress and high productivity conditions to positive under high stress and low productivity conditions. In this study, we explore the relative strength and direction of marsh plant interactions at early life stages across a stress and productivity gradient in South Slough, Coos Bay, Oregon, USA. Germination and survival of five plant species (Atriplex patula L., Distichlis spicata (L.), Plantago maritima (Lam.) A. Gray, Sarcocornia pacifica (Standl.) A. J. Scott, Triglochin maritimum L.) were examined in the presence and absence of neighboring vegetation at three intertidal levels in each of three marsh sites along an estuarine gradient. We found that many of the interactions measured across these gradients were negative or neutral. The direction of the interactions depended on life stage, with neighboring plants having mostly negative and neutral effects on seed germination, and mostly neutral and rarely positive effects on seedling survival especially at lower intertidal locations. The exception was in the high intertidal at all sites, where competition was common. We found that the intensity of the interactions varied depending on marsh site, intertidal elevation, and plant species identity and that life stage and regional climate may be responsible for the general lack of positive interactions. We suggest that measuring species interactions across complex gradients of physical stress at different life stages can help refine our conceptual models and lead to better predictions of the factors controlling community structure.

CO2 gas exchange, transpiration, stomatal conductance, water use efficiency and chlorophyll content were investigated at the leaves of three species of the family Plantaginaceae: Plantago maritima L., Plantago subpolaris Andrejev and Plantago schrenkii C.Koch under natural conditions of the habitat on high and low tide areas of the White Sea’ coasts. The high rate of photosynthesis at saturating CO2 (PNmax) at P. maritima (85.0±4.8 μmol m-2 s-1), as compared to P. subpolaris and P. schrenkii (45.2± 7.5 and 36.9±3.2 μmol m-2 s-1) was caused by high activity of ribulose-1,5-bisphoshate carboxylase/oxygenase (RuBPCO), the rate of electron transport, the rate of triose phosphate utilization, TPU), as well as high efficiency of carboxylation. The rates of photosynthesis at ambient concentration of CO2 (PN) at P. maritima were 1.4 and 1.7 times higher compared to P. subpolaris and P. schrenkii. Plants of P. schrenkii are characterized by lower values of stomatal conductance and water use efficiency compared to P. maritima and P. subpolaris. In natural habitat, the limiting factor of CO2 assimilation P. subpolaris is the rate of photosynthetic electron transport, the activity of RuBPCO in P. schrenkii.

Natural habitats on the territory of Novi Sad are almost fully destroyed today, as well as their characteristic plant taxa. The reason for disappearance of natural habitats is the development of suburban communities, which is an irreversible process. Plant taxa, specific for wet, salty, and sandy ecosystems grew on those habitats twenty years ago and earlier. This paper presents the overview of 9 taxa (Suaeda maritima subsp. maritima, Androsace elongata subsp. elongata, Cirsium boujartii subsp. boujartii, Aster sedifolius subsp. canus, Blackstonia perfoliata subsp. serotina, Plantago maritima subsp. maritima, Salvia nutans, Allium angulosum, and Typha schuttleworthii). These taxa presented integral parts of autochthonous flora of Novi Sad. Since some of these taxa were found in the field 21 years ago and some even 93 years ago, they are extinct from the flora of Novi Sad.

Most salt marshes are dominated by graminoids, but patches dominated by a diverse assemblage of perennial forbs, known as forb pannes, occur on marshes in north temperate areas. These pannes and their associated species diversity appear to be highly responsive to anthropogenic change, including climate warming. We mapped all of the forb pannes on a salt marsh in Wells, Maine in order to 1) determine if geospatial features were related to panne location and species composition, and 2) record a baseline to compare with future responses to anthropogenic change. Forb pannes occupied approximately 5% of the high marsh surface, with sizes ranging from 7 m2 to 5000 m2. Pannes themselves were clustered at river/creek edges and near pools. Within pannes, species diversity was highest away from pools and the river mouth, and relative abundance of Plantago maritima was related to spatial variables. These results suggest that forb panne distribution and community structure are influenced by spatial variables and reinforce the notion that forb pannes may be used to track salt marsh responses to global change.

A vanishing species in Poland - Plantago maritima L. was regenerated in vitro from tips of shoots (obtained in vitro) and from different explants of 4-week-old seedlings: seedling tips, hypocotyls, cotyledons, roots. Murashige and Skoog basal medium, supplemented with 0.6 pM indole-3-acetic acid in combination with cytokinins 6-benzyladenine, zeatin or kinetin, was used. The plants obtained in the result of micropropagation were normal in appearence. It was proved that Plantago maritima species was amenable to propagation from different kinds of explants. The method may be of significance for protection of sea plantain.

This paper presents an histoanatomical and ecophysiological study of Plantago maritima. Studies were conducted to assess the diversity of anatomical adaptations of vegetative organs (roots, aerial stems, rhizomes and leaves) in this taxa. Results are presented with original photographs. The analysis of leaf anatomy in P. maritima showed that the leaves contained xeromorphic traits (high development of palisade and water storage parenchyma). Stomatal density is high for both epidermis and cuticle is very thick. Analyses made in the scope of the present study indicated that total ash content of the dry leaf matter is 11.98%. Rhizome is strong and shows aeriferous tissue. Leaf relative water content (LRWC) values indicates that juvenile leaves harvested from eroded and nude soil, exposed to salt stress and heat can accumulate more water.

Medium-term (5.5 years) changes in the cover of major species in “Artemisia saline puszta” (Ass) and “Pannonic Puccinellia limosa hollow” (PPlh) grassland communities in the Kiskunsag region, Hungary, were monitored and analyzed in relation to abiotic factors (e.g. air temperature, precipitation, soil moisture, salinity and alkalinity). Soil salinity varied considerably, indicating leaching and desalinization of surface layers as the most typical process occurring in the region. Yearly average covers of Artemisia santonicum and Plantago maritima were negatively and positively related to surface soil salinity, respectively, in accordance with their salt tolerance. Multiple regression analysis showed soil pH and salinity to be the most important factors determining yearly average cover of plants at Ass. Increasing pH increased the cover of A. santonicum and P. maritima, but decreased the cover of Podospermum canum. Increasing salinity decreased the cover of A. santonicum and P. canum. At PPlh, pH of groundwater had a positive effect and the lakewater level had a negative effect on the cover of Puccinellia limosa. The results provide information on the ongoing changes in the soil properties and the resulting changes in plant composition in these Hungarian salt-affected grasslands.

1. The effects of moisture conditions, seed morphology, vegetation structure and hydrodynamic variables on seed retention were examined in a system where the dominant dispersal agent is water. Experiments were conducted in a tidal salt marsh and in a flume facility where hydrodynamic variables could be controlled.2. Moisture condition of seeds greatly influenced which factors were most important in determining seed retention. Seed type (buoyancy) was the most important factor when seeds were dry with seeds possessing very low floating capacity (Plantago maritima) being retained in greater numbers than seeds with intermediate floating capacities (Suaeda maritima and Elytrigia atherica).3. In contrast, hydrodynamic variables dominated retention processes when seeds were waterlogged. The application of waves in addition to flow velocity dislodged more seeds than flow velocity alone.4. Vegetation structure influenced retention in both dry and wet conditions but less so than other factors. Denser, less rigid vegetation types retained greater numbers of seeds than more open, more rigid vegetation types.5. Results suggest that buoyancy traits appear to determine whether seeds move in the drier summer and autumn months after initial detachment from parent plants but the intensity of wave action will determine whether waterlogged seeds stay in a microsite during the wetter months of late autumn to early spring.

In salt-marshes, bulk carbon isotope values reflect organic matter contributions not only from salt-marsh plants, but also from algal and bacterial inputs, as well as allochthonous terrestrial organic matter. In the present study, compound specific isotope analysis was applied to a core and modern plant samples collected from a Machiasport, Maine salt-marsh. We sampled 10 plant species common to the marsh including Spartina alterniflora, Spartina patens, Juncus gerardi, Solidago sempervirens, Salicornia europa, Potentilla anserina, Atriplex patula, Plantago maritima, Suaeda maritima and Limonium nashii. Isolation of two n-alkanes (C27 and C29) present in marsh plant samples allowed for the reconstruction of past plant communities present in the cored deposits. For the C27 and C29 homologues there is a 3–10‰ separation in carbon isotopic composition between C3 and C4 species. Carbon isotopic values down core show a transition from a C4 dominated plant community to a C3 dominated community, attributable to the migration of lower marsh zones onto brackish/freshwater marsh zones during a late Holocene relative sea-level rise. The two homologues used have the potential to provide paleoenvironmental information in future core work, where the sea-level variable will be controlled.

We investigated the effects of increased atmospheric CO2 on the biomass, photosynthesis, protein and phenolic concentrations and content of Plantago maritima and Armeria maritima. This enabled us to test the protein competition model (PCM) for predicting C allocation to phenolics. Three contrasting responses to elevated CO2 (600 mmol CO2 mol1) between the two study species were observed. (1) In P. maritima, plant biomass increased and the maximum carboxylation rate of Rubisco (Vc,max) was decreased. However, in A. maritima, shoot biomass decreased and the Vc,max of Rubisco was unchanged. (2) The total phenolic content increased in P. maritima but decreased in A. maritima. (3) Protein concentrations and content decreased in P. maritima and rootprotein concentrations and content increased in A. maritima. We conclude that C and N allocation to phenolics and proteins is species- and organ-specific and the PCM predictions were correct when phenolics and proteins were expressed on a per plant content basis.

Theory predicts that the sex ratio of gynodioecious populations (in which hermaphrodites and females coexist) will be affected by the relative female fitness of females and hermaphrodites, and by founder events and genetic drift in small populations. We documented the sex ratio and size of 104 populations of the gynodioecious, perennial herb Plantago maritima in four archipelagos in eastern Sweden and western Finland (from latitude 53 to 64 degrees N). The sex ratio varied significantly both among and within archipelagos (range 0-70% females, median 6.3% females). The frequency of females was highest in the northernmost archipelago and lowest in the southernmost archipelago. As predicted, females were more frequently missing from small than from large populations, and the variance in sex ratio increased with decreasing population size. The relative fecundity of female plants (mean seed output per female/mean seed output per hermaphrodite) ranged from 0.43 to 2.16 (median 1.01, n = 12 populations). Among the 12 populations sampled for seed production (four in each of three archipelagos), the frequency of females was positively related to relative fecundity of females and negatively related to population size. The results suggest that the local sex ratio is influenced both by the relative fecundity of females and hermaphrodites and by stochastic processes in small populations.

Dominant halophytes, such as Plantago maritima, Aster tripolium, Artemisia santonicum, Puccinellia limosa, Festuca pseudovina and Lepidium crassifolium were monitored for their colonization by arbuscular mycorrhizal fungi (AMF) for two vegetation periods, sampled by monthly frequency. Two saline sites (A: Apaj-puszta and Z: Zabszék) were selected for the survey due to their similar physical and chemical soil characteristics (including the salt level and salt specific anions) and the distinct site use (pasturing or not) or water regime (as drought or temporally flooded at the vicinity of the Zabszék lake). Colonization of arbuscular mycorrhizal fungi greatly depended on the plant species and their physiological status, such as the mycotroph or non-mycotroph character. In this respect the highest mycorrhizal intensity (M%) and arbusculum richness (A%) was found on the Plantago maritima , while Puccinellia limosa was the less dependent species on mycorrhiza fungi. A characteristic seasonal dynamism was found at both saline sites, for all sampled halophytes. A maximum mycorrhizal colonization was recorded in late spring and early summer, when there was an increased plant-physiological initiative for the helpful symbiosis (i.e. intensive vegetative and/or generative phases of the hosts). The rate of fungal intensity in the root system (M%), and especially the arbusculum richness (A%) on the other hand was found to be dependent on the site use and the water fluctuation in the soils. A more versatile dynamism of the mycorrhizal colonization was found therefore at the vicinity of the Zabszek lake with the permanent water fluctuations. Under more drought stressed conditions an increased mycorrhizal colonization and functioning (arbusculum richness) was found, generally preceeding the high environmental stress, which was deleterious for both symbiont partners. Such mycorrhizal dynamism in the rhizosphere, however, seems to be a common strategy between the hosts and the microsymbionts in the “struggle for life” process in the Hungarian steppe.