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Ethylene is a strong controller of root development, and it has been suggested that it is involved in the increase of lateral root development in nutrient-rich soil patches (selective root placement). Here, this contention was tested by comparing selective root placement of an ethylene-insensitive transgenic tobacco (Nicotiana tabacum) genotype (Tetr) with that of wild-type (Wt) plants. Wt and Tetr plants were grown in pots with locally increased nitrate or phosphate concentrations, after which the root growth patterns were compared with those of plants grown in pots with homogeneous nutrient distribution. Tetr plants responded to nutrient patches in a similar way to Wt plants, by placing their roots preferentially at locations with higher nutrient content, and other aspects of plant morphology were also not greatly influenced by ethylene insensitivity. The response of both Wt and Tetr plants to high-nitrate patches was considerably stronger than to locally high phosphate, suggesting that the two responses are not linked in any functional or regulatory way. As the response to nutrient patches was similar in ethylene-sensing and ethylene-insensitive plants, it is concluded that selective root placement in response to nitrate or phosphate is, at least in tobacco, not mediated or modified by ethylene action.

Flooded plant roots commonly form aerenchyma, which allows gas diffusion between shoots and roots. The programmed cell death involved in this induced aerenchyma formation is controlled by the plant hormone ethylene, as has been shown for maize (Zea mays). However, the role of ethylene is uncertain in wetland species that form constitutive aerenchyma (also under nonflooded conditions). The aim of this study is to shed light on the involvement of ethylene in constitutive aerenchyma formation in Juncus effusus. Plants of J. effusus and maize were treated with ethylene and inhibitors of ethylene action to determine the consequences for aerenchyma formation. Neither treatment with high ethylene concentrations nor with ethylene inhibitors resulted in changes in root aerenchyma in J. effusus. By contrast, ethylene increased aerenchyma development in maize unless ethylene action inhibitors were applied simultaneously. Similarly, root elongation was insensitive to ethylene treatment in J. effusus, but was affected negatively in maize. The data show that aerenchyma in J. effusus is highly constitutive and, in contrast to the inducible aerenchyma in maize, is not obviously controlled by ethylene.

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.

Terrestrial plants experience multiple stresses when they are submerged, caused both by oxygen deficiency due to reduced gas diffusion in water, and by shade due to high turbidity of the floodwater. It has been suggested that responses to submergence are de facto responses to low light intensity. We investigated the extent to which submergence and shade induce similar acclimation responses by comparing two terrestrial Rumex species that differ in their responses to flooding. Our study confirms that there are strong similarities between acclimation responses to shade and submergence. Petiole length, specific leaf area (SLA), chlorophyll parameters and underwater light-compensation points changed at least qualitatively in the same direction. Maximum underwater photosynthesis rate, however, did discriminate between the functionality of the responses, as the acclimation to submergence appeared to be more effective than acclimation to shade at saturating light. We conclude that acclimation to submergence involves more than an increase in SLA to achieve the significant reduction of diffusion resistance for gas exchange between leaves and the water column.

A plant's ability to produce and respond to ethylene is essential for its vegetative growth. We studied whole-shoot ethylene emission and leaf growth responses to applied ethylene in four Poa spp. that differ inherently in leaf elongation rate and whole-plant relative growth rate. Compared with the fast-growing Poa annua and Poa trivialis, the shoots of the slow-growing species Poa alpina and Poa compressa emitted daily 30% to 50% less ethylene, and their leaf elongation rate was more strongly inhibited when ethylene concentration was increased up to 1 microL L(-1). To our surprise, however, low ethylene concentrations (0.02-0.03 microL L(-1)) promoted leaf growth in the two slow-growing species; at the same concentrations, leaf elongation rate of the two fast-growing species was only slightly inhibited. All responses were observed within 20 min after ethylene applications. Although ethylene generally inhibits growth, our results show that in some species, it may actually stimulate growth. Moreover, in the two slow-growing Poa spp., both growth stimulation and inhibition occurred in a narrow ethylene concentration range, and this effect was associated with a much lower ethylene emission. These findings suggest that the regulation of ethylene production rates and perception of the gas may be more crucial during leaf expansion of these species under non-stressful conditions and that endogenous ethylene concentrations are not large enough to saturate leaf growth responses. In the two fast-growing species, a comparatively higher ethylene endogenous concentration may conversely be present and sufficiently high to saturate leaf elongation responses, invariably leading to growth inhibition.

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.

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.

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.