In O(2)-free media, as in wetlands, the COPR for roots is likely to be quite low, dependent upon the respiratory demands, dimensions and diffusion characteristics of the stele/stelar meristem and the enzyme kinetics of cytochrome oxidase. Roots of non-wetland plants may not differ greatly in their COPRs from those of wetland species. There is a possibility that trace amounts of O(2) may still be present in stelar 'anaerobic' cores where fermentation is induced at low cortical OPPs.
A root pressure probe and transport of
36Cl- from the medium to the xylem
were used to test whether development of oxygen deficiency in the stele, but
not the cortex, reduces radial solute transport in excised primary roots of
maize. Oxygen micro-electrodes demonstrated that a core of anoxia developed
within the stele of roots exposed to 0.05 mol m-3
O2 at 25°C, and that oxygen concentrations in the
pericycle were near or below the
Km for O2
uptake by cells, while oxygen concentrations in the endodermis, cortex and
epidermis were sufficient to fully support oxidative phosphorylation.
Decreasing the external O2 concentration from 0.27 mol
m-3 (aerated) to 0.05 mol m-3
decreased root pressure by about 45% to a new steady state over
4–6 h. Uptake of 36Cl- by
roots grown without Cl- demonstrated that the decrease
in root pressure at low O2 concentrations was almost
fully accounted for by a decrease in the rate of net radial energy-dependent
ion transport into the xylem. Even so, some energy-dependent transport
continued at low O2 supply. The data suggest that
energy-dependent solute transport into the xylem is one of the first processes
adversely affected when O2 supply to roots is low, and
provide further evidence against the classical theory of radial solute
transport (Crafts-Broyer hypothesis).
In a study on the mechanism of stimulated petiole elongation in submerged plants, oxygen concentrations in petioles of the flood-tolerant plant Rumex palustris were measured with micro-electrodes. Short-term submergence lowered petiole partial oxygen pressure to c. 19 kPa whereas prolonged submergence under continuous illumination depressed oxygen levels to c. 8-12 kPa after 24 h. Oxygen levels in petioles depended on the presence of the lamina, even in submerged conditions, and on available light. In darkness, petiole oxygen levels in submerged plants dropped quickly to values as low as 0.5-4 kPa. It is hypothesized that prolonged submergence in the light is accompanied by a decrease in carbon dioxide in the petiole. Submergence-enhanced petiolar elongation rate was compared with emergent plants. Peak daily elongation rates occurred at the end of the dark period in emergent plants, but in the middle of the light period in submerged plants. We suggest that this shift in daily elongation pattern is induced by dependence of growth on photosynthetically derived oxygen in submerged plants. Implications of reduced oxygen for ethylene production are raised. Levels of 1-aminocyclopropane-1-carboxylic acid synthase and 1-aminocyclopropane-1-carboxylic acid oxidase and ethylene sensitivity are cited as potential factors in hypoxia-induced ethylene release.
The aim of this study was to determine the relationship between shoot nitrate concentration, mediated by nitrate supply to roots, and root exudation from Hordeum vulgare. Plants were grown for 14 d in C-free sand microcosms, supplied with nutrient solution containing 2 mM nitrate. After this period, three treatments were applied for a further 14 d: (A) continued supply with 2 mM nitrate (zero boost), (B) supply with 10 mM nitrate (low boost), and (C) supply with 20 mM nitrate (high boost). At the end of the treatment period, a bacterial biosensor (Pseudomonas fluorescens 10586 pUCD607, marked with the lux CDABE genes for bioluminescence) was applied to the microcosms to report on C-substrate availability, as a consequence of root exudation. The nitrate boost treatments significantly affected shoot nitrate concentrations, in the order C>B>A. In treatments receiving a nitrate boost (B, C), increased shoot nitrate concentration was correlated with increased plant biomass, reduced root length, reduced number of root tips, and increased mean root diameter, relative to the no boost treatment (A). Imaging of biosensor bioluminescence (proportional to metabolic activity in response to availability of root exudates) indicated that root exudation increased with decreasing shoot nitrate concentration. Biosensor reporting of root C-flow indicated that exudation was greater from root tip regions than from the whole root, but that specific exudation rates for all sites were unaffected by treatments. Total root exudation across treatments was found to be closely correlated with total root length, indicating that increased root exudation, per unit root biomass, with decreasing nitrate supply was associated with altered root morphology, as a consequence of systemic plant responses to internal N-status.
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