Physiological Adaptation and Plant Distribution along a Steep Hydrological Gradient.

Plant species often separate strongly along steep environmental gradients. Our objective was to study how coupling between plant physiology and environmental conditions shapes vegetation characteristics along a distinct hydrological gradient. We therefore investigated species photosynthesis in air and under water within a limited area from dry-as-dust to complete submergence in a nutrient-poor limestone habitat on Öland's Alvar, Sweden. We found structural and physiological adaptations of species to endure water limitation at the dry end (e.g., moss cushions and CAM-metabolism) and diffusive carbon limitation (e.g., bicarbonate use) at the submerged end of the gradient. As anticipated, mean photosynthesis in air increased 18-fold from the species-poor assembly of cushion-mosses and Sedum CAM-species on mm-thin limestone pavements to the species-rich assembly of C-3 terrestrial plants in deeper and wetter soils. A GLM-model indicated that 90% of the variation in species richness could be explained by a positive effect of soil depth, a negative effect of the duration of water cover and their interaction. In water, mean photosynthesis was highest among aquatic species, low among Sedum species and cushion mosses, and negligible among C-3 terrestrial plants. While aquatic species dried out in air, drought-resistant small species were probably competitively excluded from the more suitable terrestrial habitats on deeper soils with moderate flooding by taller species of high photosynthetic capability. In conclusion, the clear distribution of species along the steep hydrological gradient reflects distinct structural and physiological adaptations, environmental filtering and interspecific competition.

Shallow plant-dominated lakes - extreme environmental variability, carbon cycling and ecological species challenges.

BACKGROUND: Submerged plants composed of charophytes (green algae) and angiosperms develop dense vegetation in small, shallow lakes and in littoral zones of large lakes. Many small, oligotrophic plant species have declined due to drainage and fertilization of lakes, while some tall, eutrophic species have increased. Although plant distribution has been thoroughly studied, the physiochemical dynamics and biological challenges in plant-dominated lakes have been grossly understudied, even though they may offer the key to species persistence. SCOPE: Small plant-dominated lakes function as natural field laboratories with eco-physiological processes in dense vegetation dictating extreme environmental variability, intensive photosynthesis and carbon cycling. Those processes can be quantified on a whole lake basis at high temporal resolution by continuously operating sensors for light, temperature, oxygen, etc. We explore this hitherto hidden world. CONCLUSIONS: Dense plant canopies attenuate light and wind-driven turbulence and generate separation between warm surface water and colder bottom waters. Daytime vertical stratification becomes particularly strong in dense charophyte vegetation, but stratification is a common feature in small, shallow lakes also without plants. Surface cooling at night induces mixing of the water column. Daytime stratification in plant stands may induce hypoxia or anoxia in dark bottom waters by respiration, while surface waters develop oxygen supersaturation by photosynthesis. Intensive photosynthesis and calcification in shallow charophyte lakes depletes dissolved inorganic carbon (DIC) in surface waters, whereas DIC is replenished by respiration and carbonate dissolution in bottom waters and returned to surface waters before sunrise. Extreme diel changes in temperature, DIC and oxygen in dense vegetation can induce extensive rhythmicity of photosynthesis and respiration and become a severe challenge to the survival of organisms. Large phosphorus pools are bound in plant tissue and carbonate precipitates. Future studies should test the importance of this phosphorus sink for ecosystem processes and competition between phytoplankton and plants.

Heat stress of two tropical seagrass species during low tides - impact on underwater net photosynthesis, dark respiration and diel in situ internal aeration.

Seagrasses grow submerged in aerated seawater but often in low O2 sediments. Elevated temperatures and low O2 are stress factors. Internal aeration was measured in two tropical seagrasses, Thalassia hemprichii and Enhalus acoroides, growing with extreme tides and diel temperature amplitudes. Temperature effects on net photosynthesis (PN ) and dark respiration (RD ) of leaves were evaluated. Daytime low tide was characterized by high pO2 (54 kPa), pH (8.8) and temperature (38°C) in shallow pools. As PN was maximum at 33°C (9.1 and 7.2 µmol O2  m(-2) s(-1) in T. hemprichii and E. acoroides, respectively), the high temperatures and reduced CO2 would have diminished PN , whereas RD increased (Q10 of 2.0-2.7) above that at 33°C (0.45 and 0.33 µmol O2  m(-2)  s(-1) , respectively). During night-time low tides, O2 declined resulting in shoot base anoxia in both species, but incoming water containing c. 20 kPa O2 relieved the anoxia. Shoots exposed to 40°C for 4 h showed recovery of PN and RD , whereas 45°C resulted in leaf damage. These seagrasses are 'living near the edge', tolerant of current diel O2 and temperature extremes, but if temperatures rise both species may be threatened in this habitat.

Photosynthetic response to globally increasing CO2 of co-occurring temperate seagrass species.

Photosynthesis of most seagrass species seems to be limited by present concentrations of dissolved inorganic carbon (DIC). Therefore, the ongoing increase in atmospheric CO2 could enhance seagrass photosynthesis and internal O2 supply, and potentially change species competition through differential responses to increasing CO2 availability among species. We used short-term photosynthetic responses of nine seagrass species from the south-west of Australia to test species-specific responses to enhanced CO2 and changes in HCO3 (-) . Net photosynthesis of all species except Zostera polychlamys were limited at pre-industrial compared to saturating CO2 levels at light saturation, suggesting that enhanced CO2 availability will enhance seagrass performance. Seven out of the nine species were efficient HCO3 (-) users through acidification of diffusive boundary layers, production of extracellular carbonic anhydrase, or uptake and internal conversion of HCO3 (-) . Species responded differently to near saturating CO2 implying that increasing atmospheric CO2 may change competition among seagrass species if co-occurring in mixed beds. Increasing CO2 availability also enhanced internal aeration in the one species assessed. We expect that future increases in atmospheric CO2 will have the strongest impact on seagrass recruits and sparsely vegetated beds, because densely vegetated seagrass beds are most often limited by light and not by inorganic carbon.

Use of sediment CO2 by submersed rooted plants.

BACKGROUND AND AIMS: Submersed plants have different strategies to overcome inorganic carbon limitation. It is generally assumed that only small rosette species (isoetids) are able to utilize the high sediment CO(2) availability. The present study examined to what extent five species of submersed freshwater plants with different morphology and growth characteristics (Lobelia dortmanna, Lilaeopsis macloviana, Ludwigia repens, Vallisneria americana and Hydrocotyle verticillata) are able to support photosynthesis supplied by uptake of CO(2) from the sediment. METHODS: Gross photosynthesis was measured in two-compartment split chambers with low inorganic carbon availability in leaf compartments and variable CO(2) availability (0 to >8 mmol L(-1)) in root compartments. Photosynthetic rates based on root-supplied CO(2) were compared with maximum rates obtained at saturating leaf CO(2) availability, and (14)C experiments were conducted for two species to localize bottlenecks for utilization of sediment CO(2). KEY RESULTS: All species except Hydrocotyle were able to use sediment CO(2), however, with variable efficiency, and with the isoetid, Lobelia, as clearly the most effective and the elodeid, Ludwigia, as the least efficient. At a water column CO(2) concentration in equilibrium with air, Lobelia, Lilaeopsis and Vallisneria covered >75% of their CO(2) requirements by sediment uptake, and sediment CO(2) contributed substantially to photosynthesis at water CO(2) concentrations up to 1000 micromol L(-1). For all species except Ludwigia, the shoot to root ratio on an areal basis was the single factor best explaining variability in the importance of sediment CO(2). For Ludwigia, diffusion barriers limited uptake or transport from roots to stems and transport from stems to leaves. CONCLUSIONS: Submersed plants other than isoetids can utilize sediment CO(2), and small and medium sized elodeids with high root to shoot area in particular may benefit substantially from uptake of sediment CO(2) in low alkaline lakes.

Contrasting oxygen dynamics in the freshwater isoetid Lobelia dortmanna and the marine seagrass Zostera marina.

BACKGROUND: and Aims Submerged plants possess well-developed aerenchyma facilitating intra-plant gas-phase diffusion of O2 to below-ground tissues, which are usually buried in anoxic sediments. However, aquatic habitats differ in terms of O2 fluctuations in the water column and in O2 consumption of the sediment, and aquatic plants differ in aerenchymal volume and resistance to O2 diffusion through the plant and across leaf and root surfaces. The hypothesis that the freshwater isoetid Lobelia dortmanna and the marine seagrass Zostera marina should display pronounced contrasts in intra-plant O2 dynamics because of differences in morphology/anatomy, physiology and growth habitat was tested. METHODS: In order to determine the O2 dynamics and relate this to the anatomy and morphology of the two species, O2 microelectrodes were inserted in the aerenchyma of leaves and roots, the sediment pore-water, and the water column in the field. Manipulation of water column O2 in the laboratory was also carried out. KEY RESULTS: It was found that intra-plant transport of O2 between leaf and root tips takes place more readily in L. dortmanna than in Z. marina due to shorter distances and greater cross-sections of the aerenchyma. The major exchange of O2 across roots of L. dortmanna can be accounted for by small intra-plant resistances to diffusion, larger root than leaf surfaces, and greater radial diffusive resistance of leaves than roots. In contrast, the major O2 exchange across leaves than roots of Z. marina can be accounted for by the opposite anatomical-morphological features. The larger aerenchymal volume and the smaller metabolic rates of L. dortmanna compared to Z. marina imply that turnover of O2 is slower in the aerenchyma of L. dortmanna and O2 fluctuations are more dampened following changes in irradiance. Also, O2 accumulated in the aerenchyma can theoretically support dark respiration for a few hours in L. dortmanna but for only a few minutes in Z. marina. CONCLUSIONS: The build-up of O2 in the pore-water of L. dortmanna sediments during the day as a result of high release of photosynthetic O2 from roots and low O2 consumption of sediments means that sediment, aerenchyma and water are important O2 sources for respiration during the following night, while Z. marina relies on the water column as the sole source of O2 because its sediments are anoxic. These differences between L. dortmanna and Z. marina appear to represent a general difference between the isoetid species mainly inhabiting sediments of low reducing capacity of oligotrophic lakes and the elodeid freshwater species and marine seagrasses mainly inhabiting sediments of higher reducing capacity in more nutrient-rich habitats.