Lukasz Kotula

Apoplastic barriers effectively block oxygen permeability across outer cell layers of rice roots under deoxygenated conditions: roles of apoplastic pores and of respiration

*Despite the importance of the barrier to oxygen losses of the roots of hygrophytes growing in wet environments devoid of oxygen, there are few data available on permeability coefficients for O(2) across outer root cell layers (P(OPR)) and how they may change in response to low O(2). *A gas perfusion technique was used to measure the P(OPR) of rice (Oryza sativa) plants grown in either aerated or deoxygenated solution. The contributions of the apoplast and of living cells to the overall P(OPR) were characterized either by blocking apoplastic pores with precipitates of brown Cu(2)[Fe(CN)(6)] or by killing cells with 0.1 N HCl. *Compared with that of plants from aerated hydroponics, the P(OPR) of plants grown in deoxygenated medium was smaller by an order of magnitude. Precipitates resulting from CuSO(4)/K(4)[Fe(CN)(6)] treatment only formed in plants grown in aerated solution, where they reduced the P(OPR) by 5-20%. Killing of root segments with HCl increased P(OPR) in plants grown in both conditions by 20-55%. *The results indicated that apoplastic barriers effectively restricted radial O(2) loss. The relative role of the respiratory O(2) consumption of root peripheral layers increased as P(OPR) decreased.

Measurements of oxygen permeability coefficients of rice (Oryza sativa L.) roots using a new perfusion technique

A new approach is described to analyse the barrier properties of the outer part of rice (Oryza sativa L.) roots towards oxygen. By using a root-sleeving O2 electrode, radial oxygen loss at different distances from the root apex was measured and related to the corresponding root structure. In addition, internal oxygen concentrations were precisely adjusted using a newly developed perfusion technique. Thus, the oxygen permeability coefficient of the outer part of the root (OPR) could be calculated, since both (i) the oxygen flow across the OPR and (ii) the oxygen concentration gradient across the OPR from inside to outside were known. On the basis of the permeability coefficient, it can be decided whether or not different rates of oxygen loss across the OPR are due to changes in the OPR structure and/or to changes in the concentration gradient. The technique was applied to rice root segments, which enabled rapid perfusion of aerenchyma. In the present study, roots of rice grown under aerobic conditions were used which should have a higher O2 permeability compared with that of plants grown in deoxygenated solution. Both radial oxygen losses and permeability coefficients decreased along the root, reaching the lowest values at the basal positions. Values of oxygen permeability coefficients of the OPR were corrected for external unstirred layers. They decreased from (2.8±0.2)×10−6 m s−1 at 30 mm to (1.1±0.2)×10−6 m s−1 at 60 mm from the apex (n=5; ±SE). They were similar to those measured previously for cuticles. Low diffusional oxygen permeability of the OPR suggested that the barrier to radial oxygen loss was effective. This may help to retain oxygen within the root and enhance diffusion of oxygen towards the apex in the presence of a relatively high water permeability. The results are discussed in terms of the inter-relationship between the water and oxygen permeabilities as roots develop in either aerated or deoxygenated (stagnant) media.

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.

Salt sensitivity in chickpea (Cicer arietinum L.): ions in reproductive tissues and yield components in contrasting genotypes

The reproductive phase in chickpea (Cicer arietinum L.) is affected by salinity, but little is known about the underlying cause. We investigated whether high concentrations of Na(+) and Cl(-) in the reproductive structures influence reproductive processes. Chickpea genotypes contrasting in tolerance were subjected to 0, 35 or 50 mm NaCl applied to soil in pots. Flower production and abortion, pod number, percentage of empty pods, seed number and size were evaluated. The concentrations of Na(+) , K(+) and Cl(-) were measured in various plant tissues and, using X-ray microanalysis, in specific cells of developing reproductive structures. Genotypic variation in reproductive success measured as seed yield in saline conditions was associated with better maintenance of flower production and higher numbers of filled pods (and thus seed number), whereas seed size decreased in all genotypes. Despite the variation in reproductive success, the accumulation of Na(+) and Cl(-) in the early reproductive tissues of developing pods did not differ between a tolerant (Genesis836) and a sensitive (Rupali) genotype. Similarly, salinity tolerance was not associated with the accumulation of salt ions in leaves at the time of reproduction or in seeds at maturity.

How belowground interactions contribute to the coexistence of mycorrhizal and non-mycorrhizal species in severely phosphorus-impoverished hyperdiverse ecosystems

BackgroundMycorrhizal strategies are very effective in enhancing plant acquisition of poorly-mobile nutrients, particularly phosphorus (P) from infertile soil. However, on very old and severely P-impoverished soils, a carboxylate-releasing and P-mobilising cluster-root strategy is more effective at acquiring this growth-limiting resource. Carboxylates are released during a period of only a few days from ephemeral cluster roots. Despite the cluster-root strategy being superior for P acquisition in such environments, these species coexist with a wide range of mycorrhizal species, raising questions about the mechanisms contributing to their coexistence.ScopeWe surmise that the coexistence of mycorrhizal and non-mycorrhizal strategies is primarily accounted for by a combination of belowground mechanisms, namely (i) facilitation of P acquisition by mycorrhizal plants from neighbouring cluster-rooted plants, and (ii) interactions between roots, pathogens and mycorrhizal fungi, which enhance the plants’ defence against pathogens. Facilitation of nutrient acquisition by cluster-rooted plants involves carboxylate exudation, making more P available for both themselves and their mycorrhizal neighbours. Belowground nutrient exchanges between carboxylate-exuding plants and mycorrhizal N2-fixing plants appear likely, but require further experimental testing to determine their nutritional and ecological relevance. Anatomical studies of roots of cluster-rooted Proteaceae species show that they do not form a complete suberised exodermis.ConclusionsThe absence of an exodermis may well be important to rapidly release carboxylates, but likely lowers root structural defences against pathogens, particularly oomycetes. Conversely, roots of mycorrhizal plants may not be as effective at acquiring P when P availability is very low, but they are better defended against pathogens, and this superior defence likely involves mycorrhizal fungi. Taken together, we are beginning to understand how an exceptionally large number of plant species and P-acquisition strategies coexist on the most severely P-impoverished soils.

Anatomical and biochemical characterisation of a barrier to radial O2 loss in adventitious roots of two contrasting Hordeum marinum accessions

A barrier to radial O2 loss (ROL) in roots is an adaptive trait of waterlogging-tolerant plants. Hordeum marinum Huds. is a waterlogging-tolerant species that, in contrast to its waterlogging-sensitive cultivated relatives, forms a tight barrier to ROL in basal root zones. To evaluate the nature of the barrier to ROL in roots, we combined measurements of ROL with histochemical and biochemical studies of two contrasting H. marinum accessions. H21 formed greater aerenchyma (up to 38% of cross-sectional area) and a tight barrier to ROL when grown under deoxygenated stagnant conditions, whereas the barrier was only partially formed in roots of H90 and aerenchyma was up to 26%. A tight barrier to ROL in roots of H21 corresponded with strong suberisation of the exodermis. In agreement with anatomical studies, almost all aliphatic suberin quantities were greater in roots of H21 grown under stagnant conditions compared with roots from aerated controls, and also to those in H90. By contrast to suberin, no differences in root lignification were observed between the two accessions raised in either aerated or stagnant conditions. These findings show that in adventitious roots of H. marinum, suberisation rather than lignification restricts ROL from the basal root zones.

Tolerance to partial and complete submergence in the forage legume Melilotus siculus: an evaluation of 15 accessions for petiole hyponastic response and gas-filled spaces, leaf hydrophobicity and gas films, and root phellem

Background and Aims Submergence is a severe stress for most plants. Melilotus siculus is a waterlogging- (i.e. root zone hypoxia) tolerant annual forage legume, but data were lacking for the effects of partial and full submergence of the shoots. The aim was to compare the tolerance to partial and full submergence of 15 M. siculus accessions and to assess variation in traits possibly contributing to tolerance. Recovery ability post-submergence was also evaluated. Methods A factorial experiment imposed treatments of water level [aerated root zone with shoots in air as controls, stagnant root zone with shoots in air, stagnant root zone with partial (75 %) or full shoot submergence] on 15 accessions, for 7 d on 4-week-old plants in a 20/15 °C day/night phytotron. Measurements included: shoot and root growth, hyponastic petiole responses, petiole gas-filled spaces, leaflet sugars, leaflet surface hydrophobicity, leaflet gas film thickness and phellem area near the base of the main root. Recovery following full submergence was also assessed. Key Results Accessions differed in shoot and root growth during partial and full shoot submergence. Traits differing among accessions and associated with tolerance were leaflet gas film thickness upon submergence, gas-filled spaces in petioles and phellem tissue area near the base of the main root. All accessions were able to re-orientate petioles towards the vertical under both partial and full submergence. Petiole extension rates were maintained during partial submergence, but decreased during full submergence. Leaflet sugars accumulated during partial submergence, but were depleted during full submergence. Growth resumption after full submergence differed among accessions and was positively correlated with the number of green leaves retained at desubmergence. Conclusions Melilotus siculus is able to tolerate partial and full submergence of at least 7 d. Leaflet surface hydrophobicity and associated gas film retention, petiole gas-filled porosity and root phellem abundance are important traits contributing to tolerance. Post-submergence recovery growth differs among accessions. The ability to retain green leaves is essential to succeed during recovery.