Radial oxygen loss and physical barriers in relation to root tissue age in species with different types of aerenchyma (original) (raw)

Root Hypoxia Reduces Leaf Growth : Role of Factors in the Transpiration Stream

PLANT PHYSIOLOGY, 1990

This study examined the potential role of restricted phloem export, or import of substances from the roots in the leaf growth response to root hypoxia. In addition, the effects of root hypoxia on abscisic acid (ABA) and zeatin riboside (ZR) levels were measured and their effects on in vitro growth determined. Imposition of root hypoxia in the dark when transpirational water flux was minimal delayed the reduction in leaf growth until the following light period. Restriction of phloem transport by stem girdling did not eliminate the hypoxia-induced reduction in leaf growth. In vitro growth of leaf discs was inhibited in the presence of xylem sap collected from hypoxic roots, and also by millimolar ABA. Disc growth was promoted by sap from aerated roots and by 0.1 micromolar ZR. The flux of both ABA and ZR was reduced in xylem sap from hypoxic roots. Leaf ABA transiently increased twofold after 24 hours of hypoxia exposure but there were no changes in leaf cytokinin levels.

Aerenchyma formation and radial O2 loss along adventitious roots of wheat with only the apical root portion exposed to O2 deficiency

Plant, Cell and Environment, 2003

This study investigated aerenchyma formation and function in adventitious roots of wheat ( Triticum aestivum L.) when only a part of the root system was exposed to O 2 deficiency. Two experimental systems were used: (1) plants in soil waterlogged at 200 mm below the surface; or (2) a nutrient solution system with only the apical region of a single root exposed to deoxygenated stagnant agar solution with the remainder of the root system in aerated nutrient solution. Porosity increased two-to three-fold along the entire length of the adventitious roots that grew into the water-saturated zone 200 mm below the soil surface, and also increased in roots that grew in the aerobic soil above the water-saturated zone. Likewise, adventitious roots with only the tips growing into deoxygenated stagnant agar solution developed aerenchyma along the entire main axis. Measurements of radial O 2 loss (ROL), taken using root-sleeving O 2 electrodes, showed this aerenchyma was functional in conducting O 2. The ROL measured near tips of intact roots in deoxygenated stagnant agar solution, while the basal part of the root remained in aerated solution, was sustained when the atmosphere around the shoot was replaced by N 2 . This illustrates the importance of O 2 diffusion into the basal regions of roots within an aerobic zone, and the subsequent longitudinal movement of O 2 within the aerenchyma, to supply O 2 to the tip growing in an O 2 deficient zone.

Formation of Aerenchyma and the Processes of Plant Ventilation in Relation to Soil Flooding and Submergence

Plant Biology, 1999

Enhanced development of gas-spaces beyond that due to the partial cell separation normally found in ground parenchymas and their derivatives creates tissue commonly termed "aerenchyma". Aerenchyma can substantially reduce internal impedance to transport of oxygen, nitrogen and various metabolically generated gases such as carbon dioxide and ethylene, especially between roots and shoots. Such transport lessens the risk of asphyxiation under soil flooding or more complete plant submergence, and promotes radial oxygen loss from roots leading to oxidative detoxification of the rhizosphere. Aerenchyma can also increase methane loss from waterlogged sediments via plants to the atmosphere. This review of the formation and functioning of aerenchyma particularly emphasises research findings since 1992 and highlights prospects for the future. Regarding formation, attention is drawn to how little is known of the regulation and processes that create schizogenous aerenchyma with its complex cell arrangements and differential cell to cell adhesion. More progress has been made in understanding lysigenous aerenchyma development. The review highlights recent work on the processes that sense oxygen deficiency and ethylene signals, subsequent transduction processes which initiate cell death, and steps in protoplast and wall degeneration that create the intercellular voids. Similarities between the programmed cell death and its causes in animals and the predictable patterns of cell death that create lysigenous aerenchyma are explored. Recent findings concerning function are addressed in terms of the diffusion aeration of roots, rhizosphere oxygenation and sediment biogeochemistry, photosynthesis and ventilation, pressurised gas-flows and greenhouse gas emissions and aspects of ventilation related to secondary thickening.

Radial Oxygen Loss from Plant Roots—Methods

Plants

In flooded soils, an efficient internal aeration system is essential for root growth and plant survival. Roots of many wetland species form barriers to restrict radial O2 loss (ROL) to the rhizosphere. The formation of such barriers greatly enhances longitudinal O2 diffusion from basal parts towards the root tip, and the barrier also impedes the entry of phytotoxic compounds produced in flooded soils into the root. Nevertheless, ROL from roots is an important source of O2 for rhizosphere oxygenation and the oxidation of toxic compounds. In this paper, we review the methodological aspects for the most widely used techniques for the qualitative visualization and quantitative determination of ROL from roots. Detailed methodological approaches, practical set-ups and examples of ROL from roots with or without barriers to ROL are included. This paper provides practical knowledge relevant to several disciplines, including plant–soil interactions, biogeochemistry and eco-physiological aspec...

Elongation by primary lateral roots and adventitious roots during conditions of hypoxia and high ethylene concentrations

Plant, Cell and Environment, 1997

Soil flooding results in unusually low oxygen concentrations and high ethylene concentrations in the roots of plants. This gas composition had a strongly negative effect on root elongation of two Rumex species. The effect of low oxygen concentrations was less severe when roots contained aerenchymatous tissues, such as in R. palustris Sm. R. thyrsiflorus Fingerh., which has little root porosity, was much more affected. Ethylene had an even stronger effect on root elongation than hypoxia, since very small concentrations (0·1 cm 3 m -3 ) reduced root extension in the two species, and higher concentrations inhibited elongation more severely than did anoxia in the culture medium. Thus, ethylene contributes strongly to the negative effects of flooding on root growth. An exception may be the highly aerenchymatous, adventitious roots of R. palustris. Aerenchyma in these roots provides a low-resistance diffusion pathway for both endogenously produced ethylene and shoot-derived oxygen. This paper shows that extension by roots of R. palustris in flooded soil depends almost completely on this shoot-derived oxygen, and that aerenchyma prevents accumulation of growth-inhibiting levels of ethylene in the root.

Root radial oxygen loss and the effects on rhizosphere microarea of two submerged plants

Polish Journal of Environmental Studies, 2015

Radial oxygen loss (ROL) has been suggested to be a major process to protect plants exposed to the anaerobic by-products of soil anaerobiosis. The aim of the present study was to test the effects of root ROL from two submerged plants (Hydrilla verticillata and Vallisneria spiralis) on the rhizosphere oxygen profile and rhizosphere microarea. Phospholipid fatty acids (PLFAs) of sediment samples were used to characterize and quantify the microbial community. The results showed clearly that there were significant differences between the two plants in radial oxygen loss, which affected rhizosphere physicochemical parameters and the microbial community. Rhizosphere total biomass, bacteria, gram-positive bacteria, actinomycetes, and microbial diversity of V. spiralis were significantly higher than those of H. verticillata. The present study highlights root ROL as a key parameter affecting the microbial community of the rhizosphere microarea.

Functional and chemical comparison of apoplastic barriers to radial oxygen loss in roots of rice (Oryza sativa L.) grown in aerated or deoxygenated solution

Journal of Experimental Botany, 2009

Radial oxygen loss (ROL) and root porosity of rice (Oryza sativa L.) plants grown in either aerated or deoxygenated (stagnant) conditions were combined for the first time with extensive histochemical and biochemical studies of the apoplastic barriers in the roots' peripheral cell layers. Growth in stagnant solution significantly affected structural and, consequently, the physiological features of rice roots. It increased adventitious root porosity by about 20% and decreased the ROL towards the base to zero at a distance of 40 mm from the apex. By contrast, roots of plants grown in aerated solutions revealed the highest rates of ROL at 30 mm from the apex. Differences in the ROL pattern along the root were related to histochemical studies, which showed an early development of Casparian bands and suberin lamellae in the exodermis, and lignified sclerenchyma cells in roots of plants grown in deoxygenated solution. In agreement with anatomical studies, absolute contents of suberin and lignin in the outer part of the roots (OPR) were higher in plants grown in deoxygenated solution. Regardless of growth conditions, the levels of suberin and lignin increased along the roots towards the base. It is concluded that radial oxygen loss can be effectively restricted by the formation of a suberized exodermis and/or lignified sclerenchyma in the OPR. However, the relative contribution of suberin and lignin in the formation of a tight barrier is unclear. Knowing the permeability coefficient across OPR for roots of plants grown in both conditions will allow a more precise understanding of the mechanisms controlling ROL.

Root aeration via aerenchymatous phellem: three-dimensional micro-imaging and radial O2 profiles in Melilotus siculus

New Phytologist, 2011

• Internal root aeration enables waterlogging-tolerant species to grow in anoxic soil. Secondary aerenchyma, in the form of aerenchymatous phellem, is of importance to root aeration in some dicotyledonous species. Little is known about this type of aerenchyma in comparison with primary aerenchyma. • Micro-computed tomography was employed to visualize, in three dimensions, the microstructure of the aerenchymatous phellem in roots of Melilotus siculus. Tissue porosity and respiration were also measured for phellem and stelar tissues. A multiscale, three-dimensional, diffusion-respiration model compared the predicted O 2 profiles in roots with those measured using O 2 microelectrodes. • Micro-computed tomography confirmed the measured high porosity of aerenchymatous phellem (44-54%) and the low porosity of stele (2-5%) A network of connected gas spaces existed in the phellem, but not within the stele. O 2 partial pressures were high in the phellem, but fell below the detection limit in the thicker upper part of the stele, consistent with the poorly connected low porosity and high respiratory demand. • The presented model integrates and validates micro-computed tomography with measured radial O 2 profiles for roots with aerenchymatous phellem, confirming the existence of nearanoxic conditions at the centre of the stele in the basal parts of the root, coupled with only hypoxic conditions towards the apex.