Seed dormancy and germination in Dodonaea viscosa (Sapindaceae) from south-western Saudi Arabia.

Dodonaea viscosa (Sapindaceae) is widespread in the mountainous highlands of the southwestern part of Kingdom of Saudi Arabia, where it is a medicinally important species for the people in Saudi Arabia. Seeds of this species were collected from Mount Atharb in Al-Baha region, at an altitude of 2100 m. The aims of this study were to determine if the seeds of D. viscosa have physical dormancy (i.e. a water-impermeable seed coat) and, if so, what treatments would break dormancy, and what conditions promote germination after dormancy has been broken. The dormancy-breaking treatments included: soaking of seeds in concentrated sulfuric acid (H2SO4) for 10 min, immersion in boiling water for 10 min and exposure to 50 °C for 1 min. After seeds had been pre-treated with H2SO4, to break dormancy, they were incubated at constant temperatures from 5 to 35 °C, under 12-h photoperiods or in continuous darkness, and germination recorded. Salinity tolerance was investigated by incubating acid-scarified seeds in different concentrations of mM NaCl in the light at 25 °C. Untreated seeds had low final germination 30%. Seeds that had been acid-scarified, immersed in boiling water or exposed to 50 °C all achieved 91% subsequently when incubated at 25 °C. Thus, seeds of this species in Saudi Arabia have physical dormancy, which can be broken by all three treatments designed to increase the permeability of the testa. After pre-treatment, there was a broad optimum constant temperature for germination that ranged between 5 and 25 °C but germination was inhibited by higher temperatures (30 and 35 °C). Light had little effect on this germination response. Scarified seeds were also sensitive to salinity, with the highest germination in distilled water and complete inhibition in 400 mM NaCl. Seeds that failed to germinate in saline treatments were mostly able to germinate on transfer to distilled water, suggesting osmotic inhibition.

Growth and photosynthetic responses of the cordgrass Spartina maritima to CO2 enrichment and salinity.

Future climatic scenarios combine increasing concentrations of atmospheric CO(2) and rising sea levels. Spartina maritima is a C(4) halophyte that is an important pioneer and ecosystem engineer in salt marshes of the Atlantic coast of southern Europe. A glasshouse experiment investigated the combined effects on its growth and photosynthetic apparatus of approximately doubling CO(2) concentration (from 380 to 700 µmol mol(-1)) at a range of salinity (0, 171 and 510 mM NaCl). We measured relative growth rates, gas exchange, chlorophyll fluorescence parameters, photosynthetic pigment concentrations, and total ash, Na(+), K(2+), Ca(2+) and N concentrations. Elevated CO(2) stimulated growth of S. maritima by c. 65% at all external salinities; this growth enhancement was associated with greater net photosynthetic rate (A) and improved leaf water relations. A increased despite a drop in stomatal conductance in response to 700 µmol mol(-1) CO(2). CO(2) and salinity had a marked overall effect on the photochemical (PSII) apparatus and the synthesis of photosynthetic pigments. Φ(PSII) values at midday decreased significantly with external salinity in plants grown at 380 µmol mol(-1) CO(2); and F(v)/F(m) and Φ(PSII) values were higher at 700 µmol mol(-1) CO(2) in presence of NaCl. Plant nutrient concentrations declined under elevated CO(2), which can be ascribed to the dilution effect caused by an increase in biomass. The results suggest that the productivity S. maritima and the ecosystem services it provides will increase in likely future climatic scenarios.

Salt stimulation of growth and photosynthesis in an extreme halophyte, Arthrocnemum macrostachyum.

Halophytes that are capable of tolerating a wide range of salinity may grow best at intermediate salinities, but the physiological mechanisms underlying positive growth responses to salinity are not clear. This work investigated the growth of Arthrocnemum macrostachyum (Moric) C. Koch (a halophytic C3 shrub) over a wide range of salinities, and the extent to which its responses can be explained by photosynthetic physiology. Growth, gas exchange and chlorophyll fluorescence characteristics of plants were examined in a glasshouse experiment; tissue concentrations of photosynthetic pigments, ash, sodium, potassium, calcium and nitrogen were also determined. Plants showed marked stimulation of growth by salt, with a broad optimum of 171-510 mm NaCl for relative growth rate (RGR). Stimulation of RGR appeared to depend mainly on an increase in specific shoot area, whereas reduced RGR at high salinity (1030 mm) could be attributed to a combination of lower unit shoot (leaf) rate and lower shoot mass fraction. The non-saline treatment plants had the greatest fraction of non-photosynthetic, atrophied surface area. However, net photosynthesis (A) was also stimulated by NaCl, with an optimum of c. 510 mm NaCl. The responses of A to salinity could be accounted for largely by limitation by stomatal conductance (Gs) and intercellular CO(2) concentration (Ci). Even the most hypersaline treatment apparently had no effect on photosystem II (PSII) function, and this resistance could be an important strategy for this halophyte in saline soils. In contrast, Fv/Fm indicated that absence of salt represents an environmental stress for A. macrostachyum and this could be a contributory factor to salt stimulation of A. Notwithstanding the importance of the ability to develop and maintain assimilatory surface area under saline conditions, stimulatory effects on A also appear to be part of a suite of halophytic adaptations in this plant.

Assessment of the risk posed by the antifouling booster biocides Irgarol 1051 and diuron to freshwater macrophytes.

Antifouling paints are used to reduce the attachment of living organisms to the submerged surfaces of ships, boats and aquatic structures, usually by the release of a biocide. Two 'booster' biocides in common use are the triazine herbicide Irgarol 1051 (N-2-methylthio-4-tert-butylamino-6-cyclopropylamino-s-triazine), and diuron (1-(3,4-dichlorophenyl)-3,3-dimethylurea), which are designed to inhibit algal photosynthesis. Previous research has been directed at the effects of these compounds in marine and estuarine environments. In 2001 we sampled the main rivers and shallow freshwater lakes (Broads) of East Anglia UK for Irgarol 1051, its metabolite GS26575 (2-methylamino-4-tert-butylamino-6-amino-s-triazine) and diuron in order to establish the baseline environmental concentrations of these compounds in freshwater systems of eastern UK and to investigate their possible effects on aquatic plants. Irgarol 1051, GS26575 and diuron were found in water samples collected from 21 locations. The highest concentrations were found in the Norfolk and Suffolk Broads in May. The rivers Great Ouse, Wissey, Bure and Yare also contained all three compounds, as did the Great Ouse Cut-off Channel. The toxicity of these biocides to three macrophyte species (Apium nodiflorum, Chara vulgaris, and Myriophyllum spicatum) was investigated. Deleterious effects on relative growth rate, the maximum quantum efficiency (Fv/Fm) of photosystem II and, for Apium, root mass production were found. C. vulgaris was generally most sensitive; growth, especially of roots, was strongly affected in A. nodiflorum; growth rate of M. spicatum was sensitive to diuron. No observed effect concentrations (NOEC) were interpolated using standard toxicological analysis. These were compared with measured environmental concentrations (MEC) to determine the ranges of risk quotients (MEC/NOEC). Both Irgarol 1051 and diuron represented significant risks to A. nodiflorum and C. vulgaris in this area.

Wave-induced hydraulic forces on submerged aquatic plants in shallow lakes.

BACKGROUND AND AIMS: Hydraulic pulling forces arising from wave action are likely to limit the presence of freshwater macrophytes in shallow lakes, particularly those with soft sediments. The aim of this study was to develop and test experimentally simple models, based on linear wave theory for deep water, to predict such forces on individual shoots. METHODS: Models were derived theoretically from the action of the vertical component of the orbital velocity of the waves on shoot size. Alternative shoot-size descriptors (plan-form area or dry mass) and alternative distributions of the shoot material along its length (cylinder or inverted cone) were examined. Models were tested experimentally in a flume that generated sinusoidal waves which lasted 1 s and were up to 0.2 m high. Hydraulic pulling forces were measured on plastic replicas of Elodea sp. and on six species of real plants with varying morphology (Ceratophyllum demersum, Chara intermedia, Elodea canadensis, Myriophyllum spicatum, Potamogeton natans and Potamogeton obtusifolius). KEY RESULTS: Measurements on the plastic replicas confirmed predicted relationships between force and wave phase, wave height and plant submergence depth. Predicted and measured forces were linearly related over all combinations of wave height and submergence depth. Measured forces on real plants were linearly related to theoretically derived predictors of the hydraulic forces (integrals of the products of the vertical orbital velocity raised to the power 1.5 and shoot size). CONCLUSIONS: The general applicability of the simplified wave equations used was confirmed. Overall, dry mass and plan-form area performed similarly well as shoot-size descriptors, as did the conical or cylindrical models of shoot distribution. The utility of the modelling approach in predicting hydraulic pulling forces from relatively simple plant and environmental measurements was validated over a wide range of forces, plant sizes and species.

Predicting the hydraulic forces on submerged macrophytes from current velocity, biomass and morphology.

Aquatic macrophytes are important in stabilising moderately eutrophic, shallow freshwater lakes in the clear-water state. The failure of macrophyte recovery in lakes with very soft, highly organic sediments that have been restored to clear water by biomanipulation (e.g. in the Norfolk Broads, UK) has suggested that the physical stability of the sediment may limit plant establishment. Hydraulic forces from water currents may be sufficient to break or remove plants. Our aim was to develop a simple model that could predict these forces from plant biomass, current velocity and plant form. We used an experimental flume to measure the hydraulic forces acting on shoots of 18 species of aquatic macrophyte of varying size and morphology. The hydraulic drag on the shoots was regressed on a theoretically derived predictor (shoot biomass × current velocity1.5). Such linear regressions proved to be highly significant for most species. The slopes of these lines represent species-specific, hydraulic roughness factors that are analogous to classical drag coefficients. Shoot architecture parameters describing leaf and shoot shape had significant effects on the hydraulic roughness factor. Leaf width and shoot stiffness individually did not have a significant influence, but in combination with shoot shape they were significant. This hydraulic model was validated for a subset of species using measurements from an independent set of shoots. When measured and predicted hydraulic forces were compared, the fit was generally very good, except for two species with morphological variations. This simple model, together with the plant-specific factors, provides a basis for predicting the hydraulic forces acting on the root systems of macrophytes under field conditions. This information should allow prediction of the physical stability of individual plants, as an aid to shallow-lake management.

Using chloroplast DNA to trace postglacial migration routes of oaks into Britain.

Postglacial migration is a major factor responsible for the patterns of genetic variation we see in natural populations. Fossil pollen data indicate that early postglacial colonists such as oak, were able to take both western and eastern migration routes into Britain. Analysis at a finer level is now permitted by the use of modern molecular techniques. A 13-bp duplication in the chloroplast tRNA(Leu1) intron occurs in natural populations of East Anglian oaks, but is not found in other parts of Britain or from mainland Europe. The distribution of this marker suggests that the mutation occurred either in southern England, or during migration from the mainland, and became fixed in a source population from which East Anglia was colonized. Planting of non-native trees for roadside boundaries and in the grounds of old houses and estates, explains the absence of the marker from some East Anglian oaks.

Native oak chloroplasts reveal an ancient divide across Europe.

Glacial refugia and postglacial migration are major factors responsible for the present patterns of genetic variation we see in natural populations. Traditionally postglacial history has been inferred from fossil data, but new molecular techniques permit historical information to be gleaned from present populations. The chloroplast tRNA(Leu1) intron contains regions which have been highly conserved over a billion years of chloroplast evolution. Surprisingly, in one of these regions which has remained invariant for all photosynthetic organisms so far studied, we have found intraspecific site polymorphism. This polymorphism occurs in two European oaks, Quercus robur and Q. petraea, indicating hybridisation and introgression between them. Two distinct chloroplast types occur and are distributed geographically as eastern and western forms suggesting that these oaks are each derived from at least two separate glacial refugia.

Ribosomal DNA variation and population differentiation in Salicornia L.

Analysis of nuclear rDNA variation within and between populations of Salicornia from the salt marsh at Stiffkey, Norfolk has detected new patterns of genetic differentiation. Individual plants may have alternative 10.5 and 12 kb repeat units. A strong association between the frequency of particular genetic variants and the environmental gradient of tidal inundation was evident. Salicornia dolichostachya Moss from sandy flats on the lowest parts of the marsh, near to the seaward edge, and S. fragilis Ball & Tutin from deep creeks were both monomorphic for the 12 kb variant; in contrast, S. pusilla J. Woods, characteristic of the highest parts of the tidal range, was monomorphic for the 10-5 kb variant. The more phenotypically variable S. europaea L. and S. ramosissima J. Woods populations that are found in large areas of lower and upper marsh, respectively, were heterogeneous for rDNA variant type. Nevertheless, the frequency of the 12 kb variant was significantly higher in plants from the lower marsh than in those from upper marsh, and in plants from low-lying pans than in those from raised interfluves; the 10.5 kb variant had the converse distribution. Variation in rDNA was not obviously associated with variation in morphology, or with variations in isozyme frequency established previously. Comparison of these results with those for populations from an extensive study, ranging from Anglesey (N. Wales) to S.W. Spain and the Gulf coast of Saudi Arabia, suggested that the rDNA variation and its association with environmental variation are complex and site-dependent. Plants from Anglesey had 10.5 and 11.5 kb variants, whereas only a 10-75 kb variant was detected in material from Spain and Saudi Arabia. rDNA variants were used as genetic markers in order to test the hypothesis that Salicornia is predominantly an inbreeder; conventional breeding experiments have been hampered by its highly specialized morphology and this represents the first direct, sensitive test of an idea that has been suggested mainly on morphological and phenological grounds. Analysis of rDNA in 38 maternal plants from Stiffkey and 2112 of their progeny provided no evidence for out-crossing.

Density and the commitment of apical meristems to clonal growth and reproduction in Hieracium pilosella.

We examined responses to population density in the commitment of apical meristems to reproduction and clonal growth in a rosette-forming, stoloniferous herb (Hieracium pilosella). Despite close physiological coupling between the evocation of the terminal inflorescence bud and the development of one or more axillary buds into stolons, the allocation of meristems was extremely plastic.Genets at the higher sowing densities showed density-dependent mortality consistent with self-thinning along a-3/2 trajectory. The probability of inflorescence evocation and associated stolon development was negatively dependent on surviving density. The proportinal distribution of primary stolons amongst genets became strikingly more unequal (expressed as the Gini coefficient) with increasing density. Clonal growth was resolved into the number of primary stolons per stoloniferous genet and the extent of stolon branching (i.e. number of apices per primary stolon); both showed strongly negative density-dependence. Reproduction, expressed as the mean number of flowering capitula per stoloniferous genet, declined 15-fold with increasing density; although theoretically expected to be unity, greater values resulted from capitulum production by attached secondary rosettes and lower values reflected the increasing abortion rate of inflorescence buds with increasing density.Both the total number of apices produced per unit area and the corresponding number of reproductive apices were maximal at intermediate surviving densities (700-1,000 m-2). The balance between reproductive and clonal growth may be expressed as the probability of an apical meristem producing a capitulum, that also peaked sharply at intermediate density. This finding does not conform with linear models that predict a shift from vegetative growth to sexual reproduction with increasing population density.

Carbon dioxide exchange in leaves of Spartina anglica Hubbard.

The gas-exchange method has been used to measure net carbon dioxide assimilation at different temperatures and irradiances in leaves of salt-treated and untreated plants of Spartina anglica. The results together with those of the CO2 compensation point and leaf anatomical data clearly indicate that Spartina anglica is a high photosynthetic capacity plant. At high temperatures and irradiances leaves of the salt-treated plants were able to maintain net photosynthesis, in contrast to the behaviour of leaves of untreated plants.The ecological significance of these results is discussed.