(Halimione) portulacoides is a halophytic, C₃ shrub. It is virtually confined to coastal salt marshes, where it often dominates the vegetation. The aim of this study was to investigate its growth responses to salinity and the extent to which these could be explained by photosynthetic physiology. The responses of young plants to salinity in the range 0–700 mol m⁻³ NaCl were investigated in a glasshouse experiment. The performance of plants was examined using classical growth analysis, measurements of gas exchange (infrared gas analysis), determination of chlorophyll fluorescence characteristics (modulated fluorimeter) and photosynthetic pigment concentrations; total ash, sodium, potassium and nitrogen concentrations, and relative water content were also determined. Plants accumulated Na⁺ approximately in proportion to external salinity. Salt stimulated growth up to an external concentration of 200 mol m⁻³ NaCl and some growth was maintained at higher salinities. The main determinant of growth response to salinity was unit leaf rate. This was itself reflected in rates of CO₂ assimilation, which were not affected by 200 mol m⁻³ but were reduced at higher salinities. Reductions in net photosynthetic rate could be accounted for largely by lower stomatal conductance and intercellular CO₂ concentration. Apart from possible effects of osmotic shock at the beginning of the experiment, salinity did not have any adverse effect on photosystem II (PSII). Neither the quantum efficiency of PSII (Φ_PSII) nor the chlorophyll fluorescence ratio (F_v/F_m) were reduced by salinity, and lower mid-day values recovered by dawn. Mid-day F_v/F_m was in fact depressed more at low external sodium concentration, by the end of the experiment. The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO₂ concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.

1 The construction of a raised dike across the coastal saltmarshes of Odiel (Huelva, south-west Spain) in 1977 divided a uniform area of low-lying sediment into two lagoons with very different drainage properties. To the east of the dike, there is rapid drainage into the estuarine channel of the Odiel and Tinto rivers, whereas to the west, the development of sand spits has impeded drainage such that standing water persists long after high tides, despite the slightly higher elevation of this area. 2 Both lagoons have been colonized by isolated clones of Spartina maritima, which have locally enhanced accretion to form domed tussocks (`nucleation'). The diameter of the tussocks was highly correlated with the elevation of their sediment surface, the larger tussocks having been the earlier colonists in a generally accreting system. Spartina in the interior of the tussocks showed reduced tiller density and vigour. 3 Only in the better-drained lagoon have the central, higher areas of the Spartina tussocks been invaded by Arthrocnemum perenne, although seed is freely available in both lagoons. Arthrocnemum formed a sprawling, dense canopy and a superficial, relatively impenetrable root system; it rapidly suppressed the remaining tillers of Spartina, eventually leaving only a fringe around the edge of the tussock. Areas invaded by Arthrocnemum were characterized by a superficial layer (10 cm) of oxidizing sediment $(E_h > + 80 mV)$. Spartina-dominated areas in both lagoons remained highly reducing, even in the surface layers ($E_h < - 150 mV$). 4 In terms of successional mechanisms, primary colonization by Spartina maritima facilitates the invasion by Arthrocnemum, which only becomes established from seed on relatively well-drained, oxidising sediments. The interaction is, however, more complicated than this: declining tiller density within the tussocks prior to invasion by Arthrocnemum is consistent with an inhibition mechanism, and the superior competitive ability of Arthrocnemum, once established, suggests that a tolerance mechanism may also operate

1 Submerged plants in shallow lakes are subject to pulling forces arising from waves, currents and grazing birds. Such forces can cause anchorage failure (mainly dislodgement of the root system) or breaking failure of the stems. Both lead to loss of fitness but uprooting is more damaging because many perennial species can replace broken shoot systems. 2 We investigated 12 abundant species (Ceratophyllum demersum, Chara sp., Eleogiton fluitans, Elodea canadensis, Myriophyllum spicatum, Najas marina, Potamogeton natans, P. obtusifolius, P. pectinatus, P. pusillus, Utricularia vulgaris and Zannichellia palustris) in 28 shallow lakes in the UK and the Netherlands. We measured the anchorage and breaking strengths of individual plants of different sizes. 3 Anchorage strength depends on the cohesive strength of the sediment and the size of the root system. The undrained shear-strength of sediments in shallow lakes varied more than 50-fold, but all were substantially weaker than terrestrial soils. Anchorage strength was modelled using the product of sediment cohesive strength and four measures of root-system size. A transformation of plan-form area (raising it to the power 2/3) that represented the hemispherical surface area of the root ball was consistently the best predictor of anchorage strength. 4 Breaking strength was a linear function of stem cross-sectional area in all species. Breaking stresses were comparable with those of marine algae and non-lignified terrestrial plants. 5 The results were used, in combination with plant allometric relationships, to predict the fates of four of the species when challenged with the largest waves likely to be encountered in a 10-year period, and the even greater forces exerted by grazing birds. We show that sediment strength and plant size determine whether plants break or uproot. A careful balance between investment in anchorage and in breakage resistance is needed to survive in the fluctuating physical environment of lakes. 6 Pulling forces experienced by aquatic plants are distinct from the mainly bending forces on more rigid land plants. We provide the first theoretical and quantitative framework for understanding their effects. Anchorage failure associated with the soft sediments of eutrophic lakes is likely to be a factor in the loss of macrophyte communities and an important factor in their restoration.