Oxygen deficiency in the rooting zone occurs with poor drainage after rain or irrigation, causing depressed growth and yield of dryland species, in contrast with native wetland vegetation that tolerates such conditions. This review examines how roots are injured by O2 deficiency and how metabolism changes during acclimation to low concentrations of O2. In the root apical meristem, cell survival is important for the future development; metabolic changes under anoxia help maintain cell survival by generating ATP anaerobically and minimizing the cytoplasmic acidosis associated with cell death. Behind the apex, where cells are fully expanded, ethylene-dependent death and lysis occurs under hypoxia to form continuous, gas-filled channels (aerenchyma) conveying O2 from the leaves. This selective sacrifice of cells may resemble programmed cell death and is distinct from cell death caused by anoxia. Evidence concerning alternative possible mechanisms of anoxia tolerance and avoidance is presented.

OXYGEN DEFICIENCY AND ROOT METABOLISM: Injury and Acclimation Under Hypoxia and Anoxia

A review is presented for prospects of germplasm improvement for waterlogging tolerance in wheat, barley and oats using a mechanistic approach based on adaptive physiological traits. In `The waterlogged environments for crop production' section, the extent of waterlogging is reviewed commencing with determination of environmental factors which may limit plant growth and development in waterlogging prone regions. This highlights that different types of waterlogging may exist, there may be large spatial and temporal variation in waterlogging, and that waterlogging may be confounded in field experiments with additional environmental factors. Environmental characterisation is therefore a key step to using mechanistic approaches for germplasm improvement for target environments, for extrapolation to other environments, and for development of screening protocols under controlled conditions that accurately reflect the field environment. In the `Information on key components required for germplasm improvement' section, the genetic diversity in wheat, barley and oats for waterlogging tolerance is confirmed. Physiological mechanisms for waterlogging tolerance are diverse and can be grouped into adaptive traits relating to (1) phenology, (2) morphology and anatomy, (3) nutrition, (4) metabolism including anaerobic catabolism and anoxia tolerance, and (5) post anoxic damage and recovery. For wheat and barley, there is some genetic diversity for waterlogging tolerance at the germination stage, however the full potential seems yet to be exploited. Varietal differences in tolerance at the germination stage often differ from tolerance at later stages of development, and this supports the view that different mechanisms of tolerance exist at the whole plant and tissue level. Limited work from genetic studies indicates a high heritability for waterlogging tolerance. It is concluded that the best opportunities for germplasm improvement are for further exploration and utilisation of genetic diversity by improving selection criteria including the use of marker assisted selection. Additional opportunities are described for increasing genetic diversity using wide hybridisations and development of transgenic plants.

Review of prospects for germplasm improvement for waterlogging tolerance in wheat, barley and oats

The response of the ascorbate-glutathione cycle was investigated in roots of young wheat (Triticum aestivum L.) seedlings that were deprived of oxygen either by subjecting them to root hypoxia or to entire plant anoxia and then re-aerated. Although higher total levels of ascorbate and glutathione were observed under hypoxia, only the total amount of ascorbate was increased under anoxia. Under both treatments a significant increase in the reduced form of ascorbate and glutathione was found, resulting in increased reduction states. Upon the onset of re-aeration the ratios started to decline rapidly, indicating oxidative stress. Hypoxia caused higher activity of ascorbate peroxidase, whereas activities of monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase were diminished or only slightly influenced. Under anoxia, activities of ascorbate peroxidase and glutathione reductase decreased significantly to 39 and 62%, respectively. However, after re-aeration of hypoxically or anoxically pretreated roots, activity of enzymes approached the control levels. This corresponds with the restoration of the high reduction state of ascorbate and glutathione within 16 to 96 h of re-aeration, depending on the previous duration of anoxia. Apparently, anoxia followed by re-aeration more severely impairs entire plant metabolism compared with hypoxia, thus leading to decreased viability.

Re-Aeration following Hypoxia or Anoxia Leads to Activation of the Antioxidative Defense System in Roots of Wheat Seedlings

Root adaptation to soil waterlogging

The present paper describes the effects of growth of roots of wheat (Triticum aestivum cv. Gamenya) in hypoxic nutrient solutions on acrenchyma formation and O2 movement from shoots to roots. Two types of roots were investigated: (1) seminal roots of 4–7-d-old seedlings, and (2) seminal and nodal roots of 10–28-d-old plants. Gas-filled porosity of seminal and nodal roots increased from 3 to 12% and from 5–7 to 11–15%, respectively, when the roots emerged in stagnant or N2-flushed solutions (0.003 mol m −3 O2) compared with growth in continuously acrated solutions (0.26 mol m −3 O2). However, neither root type increased in porosity when they were longer than 100–200 mm at the start of the exposure to these stagnant or N2-flushed treatments. A vernier microscope and cylindrical platinum-electrode were used to examine the relationship between root extension and transport of O2 from shoots to roots via the gas spaces. Measurements were made when the roots were in an anoxic medium and were dependent solely on O2 supplied from the shoots. For seminal roots of 5–7-d-old seedlings raised in stagnant solutions (90–100 mm), internal O2 transport was sufficient to support a rate of root elongation in the O2-free medium of between 0.03 and 0.17 mm h−1. When the O2 pressure around the shoots was increased from 20 to 100 kPa O2, the O2 concentrations at the walls of the expanding zone (2–7 mm from the tip) of these roots increased from 0.006 mol m−3 to between 0.04 and 0.26 mol m−3, and the rate of root extension increased five-fold. Oxygen transport to roots grown continuously in acrated solutions was considerably less than for roots raised in stagnant solutions; this difference was greater for seminal than for nodal roots. When the acrated seminal roots were longer than 100 mm and transferred to an O2-free root medium, O2 concentration became zero at the root tip causing elongation to cease. After 24 h of anoxia, none of these roots were able to resume elongation following a return to acrated solutions.

Aerenchyma formation and associated oxygen movement in seminal and nodal roots of wheat.

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis I. Anatomy and Respiration in Triticum aestivum L.

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis II. Morphology and Anatomy of Evolution Forms

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis: III. Oxygen Concentration in the Roots

Anatomy of the Root-Shoot Junction in Wheat Seedlings with Respect to Internal Oxygen Transport and Root Growth Retardation by External Oxygen Shortage

B. Erdmann

Papers

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis I. Anatomy and Respiration in Triticum aestivum L.

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis II. Morphology and Anatomy of Evolution Forms

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis: III. Oxygen Concentration in the Roots

Anatomy of the Root-Shoot Junction in Wheat Seedlings with Respect to Internal Oxygen Transport and Root Growth Retardation by External Oxygen Shortage