Ecophysiological adaptations of coastal halophytes from foredunes and salt marshes

Ecophysiological strategies of coastal halophytes from foredunes and salt marshes are discussed. A comparison is made of the factors that limit growth in salt marshes and sand dunes. In salt marshes, zonation and succession are primarily governed by variation in soil salinity, which strongly depends on inundation with seawater. Results are described of experiments which aim at separating salinity and inundation effects on growth, osmotic and mineral relations in a comparison of salt-marsh halophytes. The growth response of plants cannot simply be correlated (and causally explained) with the concentration of Na, Cl, and K in the tissues. Also, the compatible osmotic solutes proline and methylated quaternary ammonium compounds may accumulate both in species with a positive response to increased salinity and in species with a growth reduction under seawater inundation. More likely inadequate adaptation of the plants water potential with these components is partly the cause of retarded growth. Disfunctioning of the plant in this respect may be at three levels: (a) total water potential of the plant, (b) (loss) of turgor pressure potential; (c) regulation at the cellular level. The ecological importance of some factors in seawater other than sodium chloride is considered. In coastal sand dunes airborne rather than soil salinity limits plant growth, together with the effects of abrasion, sand accretion, drought and the poor nutrient status of the dune sand. Adaptations of sand-dune species to these factors may consist of: large seeds with storage tissue germinating in the dark and seedling growth enough to emerge through the accreted sand. Aerial parts must be resistant to mechanical damage (high wind speed and abrasion), possibly by a sclerophyllous and tough structure. Efficient nutrient uptake, translocation and retranslocation seem to help survive sand-dune species in a nutrient-poor rooting medium.     

The Occurrence of <Emphasis Type="Italic">p</Emphasis>-coumaric Acid and Ferulic Acid in Fossil Plant Materials and their Use as UV-proxy

The applicability of p-coumaric acid and ferulic acid concentrations or ratios in (sub)fossil plant remnant as UV-B proxies relies on various aspects, which are discussed in this paper and will be illustrated with some experimental data. A newly developed THM-micropyrolysis–gas chromatography–mass spectrometry method was tested on various spores, pollen and other plant remains, which were analysed for the presence of the UV-absorbing compounds p-coumaric acid and ferulic acid. This revealed that these supposed building-blocks of sporopollenin appear to be present in pollen of many plant species but also in moss spores. The development of this micropyrolysis method paved the way for the quantitative analysis of UV-absorbing compounds in case only a small amount of analyte is available, for example for fossil pollen and spores but also other small palynomorphs and plant fossils. The use of this technique will provide a better insight in the plant responses to UV-radiation, the chemistry of pollen and spores, their fossil counterparts and furthermore the means for a further development of a proxy for the reconstruction of past UV-B radiation.     

The combined effects of CO2 concentration and solar UV-B radiation on faba bean grown in open-top chambers

The response of faba bean seedlings to the combined effects of increased atmospheric CO2 concentrations ([CO2]) and solar UV-B irradiance was studied using open-top chambers transparent to UV-B radiation. The purpose of the study was to determine whether effects of increased [CO2] on growth and physiology are modified by the present solar UV-B fluence rate in the Netherlands. Seedlings were exposed to 350 or 700 micromoles mol-1 CO2. At both [CO2], solar UV-B irradiance was either present or reduced using polyester foil opaque to UV-B radiation. To obtain information on the time dependence of increased [CO2] and UV-B radiation effects, three harvests were performed during the experiment. CO2 enrichment resulted in increased biomass production at all harvests. At the final harvest, UV-B radiation did not affect biomass production but a significant decrease was observed after 14 d of treatment. A reduction of the UV-B fluence increased shoot length at both [CO2] throughout the experiment. UV-B radiation slightly altered biomass allocation. Plants grown at reduced levels of UV-B radiation invested less biomass in flowers and more in stem material compared to plants grown at ambient UV-B levels. CO2 enrichment resulted in a stimulation of net photosynthesis after 26 and 38 d of treatment. UV-B reduction did not alter this response. After 26 d of treatment, photosynthetic acclimation to CO2 enrichment was observed, which was probably the result of accumulation of carbohydrates in the leaves. After 38 d, photosynthetic acclimation was no longer present. The UV absorbance of methanolic leaf extracts was increased by CO2 enrichment only. Both CO2 enrichment and solar UV-B reduced the transmittance of radiation through intact attached leaves. Interaction between [CO2] and UV-B radiation was limited to UV-A transmittance of leaves. Under prevalent experimental conditions, UV-B radiation did not affect the measured physiological parameters. Most open-top chambers used for climate change research are constructed of materials which do not transmit UV-B radiation. Our results indicate that part of the 'chamber effects' on plant height often described in the literature might be explained by the absence of solar UV-B radiation in these chambers.     

Size scaling of photophysiology and growth in four freshly isolated diatom species from Ryder Bay,western Antarctic peninsula

Diatoms are one of the dominant groups in phytoplankton communities of the western Antarctic Peninsula (WAP). Although generally well‐studied, little is known about size dependent photophysiological responses in diatom bloom formation and succession. To increase this understanding, four Antarctic diatom species covering two orders of magnitude in cell size were isolated in northern Marguerite Bay (WAP). Fragilariopsis sp., Pseudo‐nitzschia cf. subcurvata, Thalassiosira cf. antarctica, and Proboscia cf. alata were acclimated to three different irradiances after which photophysiology, electron transport, carbon fixation, and growth were assessed. The small species Fragilariopsis sp., Pseudo‐nitzschia cf. subcurvata, and large species Proboscia cf. alata showed similar photoacclimation to higher irradiances with a decrease in cellular chlorophyll a and an increase in chlorophyll a specific absorption and xanthophyll cycle pigments and activity. In contrast, pigment concentrations and absorption remained unaffected by higher irradiances in the large species Thalassiosira cf. antarctica. Overall, the small species showed significantly higher growth rates compared to the large species, which was related to relatively high light harvesting capacity and electron transport rates in the smaller species. However, photophysiological responses related to photoinhibition and photoprotection and carbon fixation showed no relationship with cell size. This study supports the dominance of small diatoms at low irradiances during winter and early spring, but does not provide photophysiological evidence for the dominance of large diatoms during the phytoplankton bloom in the WAP. This suggests that other factors such as grazing and nutrient availability are likely to play a major role in diatom bloom formation.     

Plant responses to atmospheric carbon dioxide enrichment: interactions with some soil and atmospheric conditions

In general, C₃ plant species are more responsive to atmospheric carbon dioxide (CO₂) enrichment than C₄-plants. Increased relative growth rate at elevated CO₂ primarily relates to increased Net Assimilation Rate (NAR), and enhancement of net photosynthesis and reduced photorespiration. Transpiration and stomatal conductance decrease with elevated CO₂, water use efficiency and shoot water potential increase, particularly in plants grown at high soil salinity. Leaf area per plant and leaf area per leaf may increase in an early growth stage with increased CO₂, after a period of time Leaf Area Ratio (LAR) and Specific Leaf Area (SLA) generally decrease. Starch may accumulate with time in leaves grown at elevated CO₂. Plants grown under salt stress with increased (dark) respiration as a sink for photosynthates, may not show such acclimation to increased atmospheric CO₂ levels. Plant growth may be stimulated by atmospheric carbon dioxide enrichment and reduced by enhanced UV-B radiation but the limited data available on the effect of combined elevated CO₂ and ultraviolet B (280–320 nm) (UV-B) radiation allow no general conclusion. CO₂-induced increase of growth rate can be markedly modified at elevated UV-B radiation. Plant responses to elevated atmospheric CO₂ and other environmental factors such as soil salinity and UV-B tend to be species-specific, because plant species differ in sensitivity to salinity and UV-B radiation, as well as to other environmental stress factors (drought, nutrient deficiency). Therefore, the effects of joint elevated atmospheric CO₂ and increased soil salinity or elevated CO₂ and enhanced UV-B to plants are physiologically complex.     

Leaf thickness and UV-B absorbing pigments of plants in relation to an elevational gradient along the Blue Mountains,Jamaica

Terrestrial plant species vary widely in their adaptation to (increasing) solar UV-B radiation. Among the various responses of higher plants to enhanced UV-B are increasing leaf thickness and increasing concentrations of UV-B absorbing compounds. In some (UV-B resistant) plant species increased leaf thickness and UV-B absorbance may form part of mechanisms protecting plants from UV-B damage. However, in UV-B sensitive plant species leaf thickness and UV-B absorbance may increase as well with enhanced UV-B radiation. In the latter case however, this response cannot prevent plant damage and disturbance. In the present field study the relationship between these plant parameters and a natural elevational UV-B gradient on the tropical island of Jamaica was described. Four plant species of the Blue Mountain Tropical Montane Forest, occurring on open forest sites along the roadside and paths were studied along an elevational gradient. Plant species studied are Redbush (Polygonum chinense), Wild ginger (Hedychium gardneranum), John Crow Bush (Bocconia frutescens) and White clover (Trifolium repens). The elevational sites were at 800, 1000, 1200, 1400 and 1600 m above sea level. Leaf thickness was measured of leaves of intact plants around midday in the field. Leaf disks (5 mm) were sampled and extracted with a methanol/HCl mixture. UV-B absorption of these leaf extracts was measured spectrophotometrically. For all species leaves from higher elevations were thicker than those from lower elevations. In addition, the absorption of UV-B of leaf extracts increased with increasing elevations. It is assumed that the calculated gradient of the UV-B_BE from 800 m above sea level: 9.45 kJ m^-2 day^-1 to 9.75 kJ m^-2 day^-1 at 1600 m is related to the measured increase of leaf thickness and UV-B absorbing compounds. The responsiveness of these plant parameters to the elevational gradient does not necessarily imply that the plant species are UV-B resistant. One possibility is that the species studied, which are growing on open, disturbed sites on the forest floor and along mountain-roads, are relatively sensitive to UV-B. In addition to clear sky conditions, mist and clouds occur frequently in this tropical mountane forest at Jamaica. Also, the low nutrient status of the soil (low pH, nutrient deficiency) and the high content of polyphenols in leaves of many plant species of the tropical montane rain forest may relate to the marked response of the species studied with increasing elevation. Abbreviations: asl – above sealevel, UV-B – ultraviolet-B radiation (280–320 nm), TMCF – Tropical Montane Cloud Forest.     

The effect of enhanced ultraviolet-B radiation on germination and seedling development of plant species occurring in a dune grassland ecosystem

The germination of seeds of seven plant species occurring in a dune grassland vegetation of the Netherlands, was studied at four levels of UV-B radiation simulating unto 45% stratospheric ozone reduction during April. With the exception of seeds of Senecio jacobaea, germination of the dune grassland species was not affected by enhanced UV-B irradiance. Although a clear UV-B fluence-response relationship was not observed, the germination rate of S. jacobaea seeds and maximal germination percentage were reduced at enhanced UV-B. Germination rate in the dark was higher than germination in the light for Oenothera biennis, Plantago lanceolata, Rumex obtusifolius and S. jacobaea. Total dry biomass accumulation of seedlings was not affected by increased UV-B radiation in any of the species tested. Clear-cut differences in UV-absorbance of methanolic extracts were observed between species. Enhanced UV-B irradiance stimulated UV-absorbance of seedling extracts of Holcus lanatus and Verbascum thapsus. A clear UV-B fluence-response relationship was observed for both species. The results indicate that germination of the studied plant species probably will not be adversely affected by the expected stratospheric ozone reduction in The Netherlands.     

The effect of solar UV radiation on four plant species occurring in a coastal grassland vegetation in The Netherlands

During the summer of 1992, growth and some physiological parameters of four native plant species occurring in a coastal grassland in The Netherlands, were studied after reduction of solar UV irradiance using different cut-off filters. Biomass production, morphology and photosynthesis of all species tested were unaffected by the different treatments. Litter production of Plantago lanceolata was increased in the absence of the total UV waveband, indicating a possible role for this waveband in plant senescence. Depletion of the total UV waveband from sunlight resulted in alterations in biomass allocation in Calamagrostis epigeios and Urtica dioica while no changes were observed in P. Ianceolata and Verbascum thapsus. In C. epigeios an increase in the specific leaf area was observed, whereas in U. dioica root weight per total plant weight was decreased resulting in an increase in the shoot/root ratio. Both photosynthetic and UV-absorbing pigment concentrations were altered by the different filter applications. When compared to control plants receiving full sunlight, depletion of UV-B resulted in a significant increase in chlorophyll concentration in U. dioica leaves, this however did not affect photosynthetic rate. The presence of UV-B radiation enhanced the UV-absorbance of leaf extracts of all species except P. lanceolata. Optical characteristics of the leaves were also changed. Both the quantity ( P. lanceolata and U. dioica ) and the quality (all species) of radiation transmitted by the leaves was affected by the different treatments.     

Effects of UV-B radiation on a charophycean alga,Chara aspera

The charophycean algal species Chara aspera was exposed for 73 days to three levels of UV-B radiation (weighted according to Caldwell's generalized plant action spectrum): 1.9 kJ m^–2 day^–1 (`no UV-B'), 6.4 KJ m<sup>–2</sup> day<sup>–1</sup> (ambient) and 10.5 kJ m<sup>–2</sup> day<sup>–1</sup> (enhanced UV-B), the latter level simulating 30% ozone reduction in The Netherlands.Charophycean algae are mainly freshwater organisms and are thought to be the algae most closely related to higher land plants. Therefore we expected that responses of charophycean algae to UV-B radiation might be more related to those observed in the higher land plants than those of other `lower' algal groups.Under elevated UV-B radiation algal length was reduced. There was no induction of UV absorbing compounds under enhanced UV-B. This might relate to a sensitive response to UV-B radiation. The charophycean algae show similar adaptations to UV-B radiation as terrestrial plants, while not having UV-screens as occur in many angiosperms. Vegetative reproduction (bulbils) increased in the presence of UV-B radiation, while generative reproduction (antheridia and oogonia) decreased.     

Consequences of depletion of stratospheric ozone for terrestrial Antarctic ecosystems: the response of Deschampsia antarctica to enhanced UV-B radiation in a controlled environment

Mini UV lamps were installed over antarctic plants at Léonie Island, Antarctic peninsula, and shoot length measurements of Deschampsia antarctica were performed during the austral summer January–February 1999.We studied the response of the antarctic hairgrass, Deschampsia antarctica to enhanced UV-B. In a climate room experiment we exposed tillers of Deschampsia antarctica, collected at Léonie Island, Antarctic peninsula, to ambient and enhanced levels of UV-B radiation. In this climate room experiment with 0, 2.5 and 5 kJ m^–2 day^–1 UV-B_BE treatments we observed that length growth of shoots at 2.5 and 5 kJ m^–2 day^–1 UV-B_BE was markedly reduced compared to 0 kJ m^–2 day^–1 UV-B_BE. In addition, there was an increased number of shoots and increased leaf thickness with enhanced UV-B. The Relative Growth Rate (RGR) was not affected by UV-B, possibly because reduced shoot length growth by enhanced UV-B was compensated by increased tillering. Light response curves of net leaf photosynthesis of plants exposed to 5 kJ m^–2 day^–1 UV-B_BE did not differ from those exposed to 0 kJ m^–2 day^–1 UV-B_BE. The content of UV-B absorbing compounds of plants exposed to increasing UV-B did not significantly change.Mini UV-B lamp systems were installed in the field, to expose the terrestrial antarctic vegetation at Léonie Island to enhanced solar UV-B. In that study, the increment of shoot length of tagged plants of Deschampsia antarctica during the January-February 1999 at Léonie Island, was recorded and compared to shoot length growth under controlled conditions.The consequences of enhanced UV-B radiation as a result of ozone depletion for the terrestrial antarctic ecosytems are discussed.