T. Luque

Growth and photosynthetic responses to copper stress of an invasive cordgrass, Spartina densiflora.

Spartina densiflora Brongn. is found in coastal marshes of south-west Spain, growing in sediments with between 300 and 3000mg Cu kg(-1) total soil DW (450-4500mg Cu kg(-1) supposing that the soil porosity is 0.5). An experiment was designed to investigate the effect of copper from 0 to 5000mgkg(-1) (64mmoll(-1)) on the photosynthetic apparatus and the growth of S. densiflora. We also determined total ash, copper, calcium, magnesium and phosphorous concentrations, as well as C/N ratio. S. densiflora survived to concentrations as high as 320mg Cukg(-1) DW in leaves, although excess of Cu diminished water use efficiency and Ca-, Mg- and P-uptake. Also, quantum efficiency of PSII, net photosynthetic rate, stomatal conductance and pigment concentrations declined with increasing external Cu. Finally, the decline in the photosynthetic function resulted in a biomass reduction of between 50 and 80% (for 600 and 5000mg Cu kg(-1), respectively).

Zinc tolerance and accumulation in the salt-marsh shrub Halimione portulacoides.

The halophytic shrub Halimione portulacoides is known to be capable of growth in soils containing extremely high concentrations of Zn. This study evaluated in detail the tolerance and accumulation potential of H. portulacoides under moderate and high external Zn levels. A greenhouse experiment was conducted in order to investigate the effects of a range of Zn concentrations (0-130 mmol L(-1)) on growth and photosynthetic performance by measuring relative growth rate, total leaf area, specific leaf area, gas exchange, chlorophyll fluorescence parameters and photosynthetic pigment concentrations. We also determined the total zinc, nitrogen, phosphorus, calcium, magnesium, sodium, potassium, iron and copper concentrations in the plant tissues. H. portulacoides demonstrated hypertolerance to Zn stress, since it survived with leaf concentrations of up to 2300 mg Zn kg(-1)dry mass, when treated with 130 mmol Zn L(-1). Zinc concentrations greater than 70 mmol L(-1) in the nutrient solution negatively affected plant growth, in all probability due to the recorded decline in net photosynthesis rate. Our results indicate that the Zn-induced decline in the photosynthetic function of H. portulacoides may be attributed to the adverse effect of the high concentration of the metal on photosynthetic electron transport. Growth parameters were virtually unaffected by leaf tissue concentrations as high as 1500 mg Zn kg(-1)dry mass, demonstrating the strong capability of H. portulacoides to protect itself against toxic Zn concentrations. The results of our study indicate that this salt-marsh shrub may represent a valuable tool in the restoration of Zn-polluted areas.

Carry-over of Differential Salt Tolerance in Plants Grown from Dimorphic Seeds of Suaeda splendens

BACKGROUND AND AIMS Halophytic species often show seed dimorphism, where seed morphs produced by a single individual may differ in germination characteristics. Particular morphs are adapted to different windows of opportunity for germination in the seasonally fluctuating and heterogeneous salt-marsh environment. The possibility that plants derived from the two morphs may also differ physiologically has not been investigated previously. METHODS Experiments were designed to investigate the germination characteristics of black and brown seed morphs of Suaeda splendens, an annual, C(4) shrub of non-tidal, saline steppes. The resulting seedlings were transferred to hydroponic culture to investigate their growth and photosynthetic (PSII photochemistry and gas exchange) responses to salinity. KEY RESULTS Black seeds germinated at low salinity but were particularly sensitive to increasing salt concentrations, and strongly inhibited by light. Brown seeds were unaffected by light, able to germinate at higher salinities and generally germinated more rapidly. Ungerminated black seeds maintained viability for longer than brown ones, particularly at high salinity. Seedlings derived from both seed morphs grew well at high salinity (400 mol m(-3) NaCl). However, seedlings derived from brown seeds performed poorly at low salinity, as reflected in relative growth rate, numbers of branches produced, F(v)/F(m) and net rate of CO(2) assimilation. CONCLUSIONS The seeds most likely to germinate at high salinity in the Mediterranean summer (brown ones) retain a requirement for higher salinity as seedlings that might be of adaptive value. On the other hand, black seeds, which are likely to delay germination until lower salinity prevails, produce seedlings that are less sensitive to salinity. It is not clear why performance at low salinity, later in the life cycle, might have been sacrificed by the brown seeds, to achieve higher fitness at the germination stage under high salinity. Analyses of adaptive syndromes associated with seed dimorphism may need to take account of differences over the entire life cycle, rather than just at the germination stage.

Short-term responses to salinity of an invasive cordgrass

Salinity is one of the main chemical factors in salt marshes. Studies focused on the analysis of salinity tolerance of salt marsh plants are very important, since they may help to relate their physiological tolerances with distribution limits in the field. Spartina densiflora is a South America cordgrass, which has started its invasion of the European coastline from the southwestern Iberian Peninsula. In this work, short-term responses in adult tussocks of S. densiflora from southwestern Spain are studied over a wide range of salinity in a greenhouse experiment. Our results point out that S. densiflora has a high tolerance to salinity, showing high growth and net photosynthesis rates from 0.5 to 20 ppt. S. densiflora showed at the lowest salinity (0.5 ppt) high levels of photoinhibition, compensated by higher levels of energy transmission between photosystems. Adaptative mechanisms, as those described previously, would allow it to live in fresh water environments. At the highest salinity (40 ppt), S. densiflora showed a high stress level, reflected in significant decreases in growth, net photosynthesis rate and photochemical efficiency of Photosystem II. These responses support S. densiflora invasion patterns in European estuaries, with low expansion rates along the coastline and faster colonization of brackish marshes and river banks.

Effects of Salinity on Germination and Seedling Establishment of Endangered Limonium emarginatum (Willd.) O. Kuntze

Abstract The germination and subsequent seedling establishment of Limonium emarginatum, an endangered and endemic halophyte of the Strait of Gibraltar, was studied under exposure to different NaCl concentrations (0, 2, 4, and 6%) in a laboratory experiment. We assessed final germination percentage, number of days to first and final germination, mean time to germinate (MTG), as well as seed viability and seedling survival. Increasing salinity delayed the beginning and ending of germination and reduced final germination percentage, inhibiting germination completely above 2% salinity. L. emarginatum exhibited the greatest germination in fresh water. When seeds were removed from all saline solutions, between 60% and 70% of final germination was recorded, although at hypersalinity, germination viability diminished. Salinity pretreatments had a stimulatory effect on germination since germination speed was higher for the recovery experiment than for the seed germination experiment. Transition between germination and seedling establishment was a critical phase, given that less than 50% of seedlings of L. emarginatum survived in distilled water and 5% survived at 2% salinity.

Chlorophyll fluorescence, stress and survival in populations of Mediterranean grassland species

. Photosynthetic stress in response to a natural episode of frost and seasonal drought was assessed in a ‘dehesa’ grassland of SW Spain with a portable fluorimeter. Chlorophyll fluorescence characteristics of dark-adapted leaves of 11 abundant species of Mediterranean grassland were measured over the course of a growing season from November 1992 to July 1993. Concomitant changes in population size were estimated from censuses of permanent quadrats. There was a general decline in the photochemical efficiency (Fv/Fm) during the growing season and this was particularly evident late in the growing season (spring and early summer) when ambient temperatures were increasing rapidly and rainfall was low; it coincided with the period of most intense mortality for most species. A frost in early March (- 5 °C), when photosynthetically active radiation was relatively high, resulted in a small decrease in Fv/Fm that was consistent across many species. The mechanisms of protection in species of Mediterranean grassland appear to be sufficiently effective to avoid damage to PSII for most of the year. For most species there was little evidence of photosystem II damage, as initial fluorescence (F0) usually did not increase. Many of the effects observed were due to a reduction in Fm and thus were consistent with non-photochemical quenching. This could be adaptive in protecting PSII from damage in species that show little evidence of stress. The sharp increase in stress toward the end of the life cycle coincided with the fall in net population size.

The role of two Spartina species in phytostabilization and bioaccumulation of Co, Cr, and Ni in the Tinto-Odiel estuary (SW Spain)

Vascular plants in salt marshes strongly influence processes of heavy metal accumulation. Many studies have focused on this issue; however, there is a lack of information regarding the effects of plants on the distribution of certain poorly studied metals, such as Co, Cr, and Ni. The aim of this study was to comparatively evaluate the capability of Spartina densiflora Brongn. and Spartina maritima (Curtis) Fernald, to accumulate Co, Cr, and Ni and influence the sediment composition around their roots, investigating whether the observed behavior can change with different levels of sediment pollution. Concentrations of Co, Cr, and Ni were determined in tissues of S. densiflora and S. maritima and in sediments and rhizosediments from the Odiel and Tinto marshes (SW Spain), one of the estuaries most polluted by heavy metals in the world. Concentrations of Co, Cr, and Ni in the belowground tissues of both Spartina species were higher than those in aboveground tissues in all sites sampled. Both species showed potential for phytostabilization of Co, possibly by promoting the formation of high amounts of Fe-oxides in the rhizosphere, which can act to retain the metal within the sediment around the roots. In addition, both Spartina species were found to accumulate Co in their roots, thereby avoiding the translocation of this metal to photosynthetic tissues. At the Tinto marsh, there were no differences recorded in metal levels between sediments and rhizosediments of both species, a fact that could be explained by the extremely high background levels of metals at this site, which may impair the ability of the plant to alter the chemistry of the sediment in contact with the roots. The potential for the immobilization of a large amount of Co in the soil, exhibited by S. densiflora and S. maritima, indicates that both species could be highly useful in the phytostabilization of Co contaminated environments.

Environmental determination of shoot height in populations of the cordgrass Spartina maritima.

Spartina species tend to exhibit a range of phenotypes, often with short and tall growth forms. Such differences have been attributed variously to environmentally induced phenotypic plasticity and genetic diferentiation between populations. This work examines the basis of height variation inSpartina maritima (Curtis) Fernald at Odiel salt marshes, southwest Spain. Populations from sites with lower sediment redox potentials tended to have significantly taller shoots. Thirty-four natural populations with an 8-fold range of shoot height were transplanted to a common environment on an unvegetated, intertidal plain and shoot height was measured annually for 3 yr. There was a striking convergence in height across populations after transplantation and the change in height in each year of a population was linearly related to its initial height. Most populations grew taller after transplantation, suggesting environmental limitation in their natural habitats. Populations that were originally tall tended to become shorter. The change in shoot height was negatively related to the difference in surface sediment redox potential between their natural sites and the common transplant site. Hypoxic sediments may stimulate stem growth, resulting in improved photosynthetic gas exchange and internal aeration of roots and rhizomes. Although height variation inS. maritima appears mainly to be a result of phenotypic plasticity, a genetic component cannot be ruled out. This study emphasizes the importance of long-term studies, preferably longer than turnover time of shoot populations. The highly plastic growth form ofS. maritima allows it to colonize a wide range of habitats in environmentally heterogeneous salt marshes.

Tolerance and accumulation of copper in the salt-marsh shrub Halimione portulacoides.

The present study evaluated the tolerance and accumulation potential in the salt-marsh shrub Halimione portulacoides under moderate and high external Cu levels. A greenhouse experiment was conducted in order to investigate the effects of a range of external Cu concentrations (0 to 60 mmol l(-1)) on growth and photosynthetic performance by measuring gas exchange, chlorophyll fluorescence parameters and photosynthetic pigments. We also determined total copper, nitrogen, phosphorus and sulfur concentrations in the plant tissues. H. portulacoides survived with external Cu concentrations of up to 35 mmol Cu l(-1), although the excess of metal resulted in a biomass reduction of 48%. The effects of Cu on growth were linked to a drastic reduction in net photosynthesis. However, H. portulacoides tolerated Cu levels of up to 15 mmol Cu l(-1) without suffering adverse physiological effects. Our results indicate that this species could play an important role in the restoration of Cu-contaminated soils.

Comparative Field Summer Stress of Three Tree Species Co-occurring in Mediterranean Coastal Dunes

Chlorophyll a fluorescence, water potential (Ψs), and root system of Juniperus oxycedrus ssp. macrocarpa, Juniperus phoenicea ssp. turbinata, and Pinus pinea were studied in Mediterranean coastal dunes of SW Spain during summer drought and after fall rains in 1999, the driest year in the 90s. A strong and reversible depression in the photochemical efficiency of photosystem 2 of the three species was recorded, which happened concomitantly with the diurnal increase and decrease in radiation. J. phoenicea, with superficial root system, was the most affected species by summer drought. It showed high rates of down-regulation of photosynthesis by photoinhibition and positive correlation between Ψs and Fv/Fp, with Ψs lower than -7 MPa. However, it tolerated this high stress, showing a fast recovery of its physiological state after fall rains. On the other hand, J. oxycedrus and P. pinea, both with deep root systems, kept their Ψs values up to -3 MPa, showing lower stress during summer drought. On the other hand, J. oxycedrus and J. phoenicea were more sensible to changes in edaphic water content than P. pinea. These specific responses to summer drought would be determined by their root distributions and stomatal control of transpiration, conditioning the efficiency in getting and using the available water resources. Ecophysiological responses indicate that these species are well-adapted to long periods of drought in Mediterranean climate areas, developing different strategies: J. phoenicea tolerates high stress with a fast recovery after fall rains, while J. oxycedrus and P. pinea are less affected by summer drought since their deep root systems would allow them to reach deep water resources.