Limited costs of wrong root placement in Rumex palustris in heterogeneous soils.

Nutrient hot spots in the soil have a limited life span, but the costs and benefits for root foraging are still underexposed. We assessed short-term costs that may arise when a nutrient-rich patch induces root proliferation, but then rapidly disappears. Rumex palustris plants were grown with a homogeneous or a heterogeneous nutrient application. After root proliferation in a nutrient-rich patch, nutrient supply was switched from homogeneous to heterogeneous, and vice versa, or the patch location was changed. R. palustris proliferated its roots in the rich patch. After switching, the relative growth rates of the roots were adjusted to the novel pattern of nutrient availability. However, the changes in local root biomass lagged behind the rapid shift in nutrient supply, because the root mass realized in specific sectors could not be rapidly relocated. Despite this, R. palustris did not exhibit costs of switching in terms of biomass or nitrogen uptake. Our data suggest that rapid shifts in uptake rate and redistribution of nitrogen within the plant may have lowered the costs of incorrect root placement.

Flooding tolerance of Carex species in relation to field distribution and aerenchyma formation.

The flooding tolerance of Carex species was studied in relation to their field distribution and their capacity to form root aerenchyma under controlled conditions. In an alpine meadow, six Carex species were selected which were distributed in a clear zonation correlating with water content of the soil. Carex sempervirens and C. ferruginea were only found on nonflooded soil, the latter species preferring moister conditions. Carex davalliana and C. nigra were both associated with water-saturated soil, whereas C. limosa and C. rostrata preferred partially submerged conditions. Carex davalliana and C. limosa were bound to flooded soils with a relatively high redox potential and horizontally flowing groundwater. Carex rostrata and C. nigra grew in stagnant soil-flooded conditions with low soil redox potentials. The amount of aerenchyma in the roots of all species increased when grown in oxygen-deficient stagnant agar. This increase in root porosity, combined with increased root diameter, presumably improved internal aeration of the roots. Although all species survived experimental soil flooding, partial submergence was lethal to C. sempervirens and, surprisingly, also to the wetland species C. davalliana. Carex ferruginea showed a reduced growth rate during partial submergence. The three other species, all wetland plants, reached highest biomass production under soil-flooded and partially submerged conditions, with slower growth on free-draining soil. It is concluded that aerenchyma is not constitutive in the Carex species under study, and is best developed in Carex species from wetlands. Species with less aerenchyma perform poorly when soil-flooded, but conditions of partial submergence could even affect species with a considerable amount of root aerenchyma.

Survival tactics of Ranunculus species in river floodplains.

The flooding resistance of four Ranunculus species was studied under controlled conditions and related to the tactics used by these species to survive in their natural habitat in river floodplains. R. bulbosus, a species from seldom-flooded river levées, was relatively intolerant of both waterlogging and complete submergence, due to a constitutively low level of aerenchyma in the root system. This lack of gas spaces resulted in high mortality rates during flooding treatments and an inability to use photosynthetically derived oxygen for root respiration during complete submergence. The pioneer R. sceleratus, predominantly abundant in low lying mudflats, was very resistant to waterlogging and shallow floods. Due to its constitutively high root porosity and its ability to greatly increase the elongation rate of petioles under water this species can ameliorate flooding stress. However, when leaf blades of R. sceleratus were unable to reach the water surface, this species died as quickly as the flooding-intolerant R. bulbosus. This indicates that fast elongation of petioles under water competes for energy and respirable reserves with maintenance processes. R. repens, a species from lower, frequently inundated floodplains, was very tolerant of prolonged waterlogging and submergence. Its high resistance to complete submergence under continuous darkness indicates that this species tolerates hypoxic and/or anoxic tissue conditions via metabolic adjustments. Lysigenous aerenchyma was also induced in the primary root system and in newly developed laterals, and it was able to use oxygen generated by underwater photosynthesis, for root respiration. R. acris, a species from less frequently flooded areas, was as resistant to waterlogging and submergence in the light as R. repens. However, it has a lower resistance than R. repens to complete submergence in the dark. A submergence pre-treatment increased the maximum net underwater photosynthetic rate in R. bulbosus, whereas a significant decrease of light compensation points was observed in R. repens when it had previously been submerged. This study shows that Ranunculus species exhibit various strategies to cope with different flooding conditions. R. repens responds to flooding by its tolerance mechanism and R. sceleratus by avoidance. R. acris ameliorates submergence and R. bulbosus was not able to adapt high water tables.

Ethylene Sensitivity and Response Sensor Expression in Petioles of Rumex Species at Low O2 and High CO2 Concentrations.

Rumex palustris, a flooding-tolerant plant, elongates its petioles in response to complete submergence. This response can be partly mimicked by enhanced ethylene levels and low O2 concentrations. High levels of CO2 do not markedly affect petiole elongation in R. palustris. Experiments with ethylene synthesis and action inhibitors demonstrate that treatment with low O2 concentrations enhances petiole extension by shifting sensitivity to ethylene without changing the rate of ethylene production. The expression level of the R. palustris gene coding for the putative ethylene receptor (RP-ERS1) is up-regulated by 3% O2 and increases after 20 min of exposure to a low concentration of O2, thus preceding the first significant increase in elongation observable after 40 to 50 min. In the flooding-sensitive species Rumex acetosa, submergence results in a different response pattern: petiole growth of the submerged plants is the same as for control plants. Exposure of R. acetosa to enhanced ethylene levels strongly inhibits petiole growth. This inhibitory effect of ethylene on R. acetosa can be reduced by both low levels of O2 and/or high concentrations of CO2.

Ethylene production and petiole growth in rumex plants induced by soil waterlogging: the application of a continuous flow system and a laser driven intracavity photoacoustic detection system.

Petiole growth of Rumex acetosa L., Rumex crispus L., and Rumex palustris Sm. in response to soil waterlogging was studied in relation to production of the gaseous plant hormone ethylene. Ethylene production was monitored in a flow-through system and a recently developed laser driven photoacoustic detection system, which allowed ethylene measurements as low as 6 picoliters per liter. R. acetosa showed a two-fold increase in ethylene production correlated with a slight enhancement of the growth of the petiole that developed during the waterlogging treatment. Both R. crispus and R. palustris showed a strong petiole elongation of existing as well as newly formed petioles, which was correlated with a 20-fold increase in ethylene production after approximately 7 days. Increased rates of ethylene production in R. palustris were related to a strong increase in 1-aminocyclopropane-1-carboxylic acid (ACC) concentration and a slight, but detectable, increase in ethylene forming enzyme activity. In R. acetosa on the other hand, only a very small increase in ACC concentration was observed. Changes in ethylene production in Rumex are strongly correlated with variation in ACC content and ethylene forming enzyme activity. The interaction between ethylene production/internal concentration and ethylene sensitivity of the three Rumex species is discussed in relation to their field location in a flooding gradient and their differential resistance toward waterlogging and submergence.