Root aeration in rice (Oryza sativa): evaluation of oxygen, carbon dioxide, and ethylene as possible regulators of root acclimatizations

Adventitious roots of rice (Oryza sativa) acclimatize to root-zone O(2) deficiency by increasing porosity, and induction of a barrier to radial O(2) loss (ROL) in basal zones, to enhance longitudinal O(2) diffusion towards the root tip. Changes in root-zone gas composition that might induce these acclimatizations, namely low O(2), elevated ethylene, ethylene-low O(2) interactions, and high CO(2), were evaluated in hydroponic experiments. Neither low O(2) (0 or 0.028 mol m(-3) O(2)), ethylene (0.2 or 2.0 microl l(-1)), or combinations of these treatments, induced the barrier to ROL. This lack of induction of the barrier to ROL was despite a positive response of aerenchyma formation to low O(2) and elevated ethylene. Carbon dioxide at 10 kPa had no effect on root porosity, the barrier to ROL, or on growth. Our findings that ethylene does not induce the barrier to ROL in roots of rice, even though it can enhance aerenchyma formation, shows that these two acclimatizations for improved root aeration are differentially regulated.     

Oxygen dynamics during submergence in the halophytic stem succulent Halosarcia pergranulata

This study elucidated O2 dynamics in shoots and roots of submerged Halosarcia pergranulata (Salicornioideae), a perennial halophytic stem succulent that grows on floodprone mudflats of salt lakes. Oxygen within shoots and roots was measured using microelectrodes, for plants when waterlogged or completely submerged, with shoots in light or in darkness, in a controlled environment. Net photosynthesis (PN) when underwater, at a range of dissolved CO2 concentrations, was measured by monitoring O2 production rates by excised stems. The bulky nature and apparently low volume of gas-filled spaces of the succulent stems resulted in relatively high radial resistance to gas diffusion. At ambient CO2, quasi-steady state rates of PN by excised succulent stems were estimated to be close to zero; nevertheless, in intact plants, underwater photosynthesis provided O2 to tissues and led to radial O2 loss (ROL) from the roots, at least during the first several hours (the time period measured) after submergence or when light periods followed darkness. The influence of light on tissue O2 dynamics was confirmed in an experiment on a submerged plant in a salt lake in south-western Australia. In the late afternoon, partial pressure of O2 (pO2) in the succulent stem was 23.2 kPa (i.e. approximately 10% above that in the air), while in the roots, it was 6.2-9.8 kPa. Upon sunset, the pO2 in the succulent stems declined within 1 h to below detection, but then showed some fluctuations with the pO2 increasing to at most 2.5 kPa during the night. At night, pO2 in the roots remained higher than in the succulent stems, especially for a root with the basal portion in the floodwater. At sunrise, the pO2 increased in the succulent stems within minutes. In the roots, changes in the pO2 lagged behind those in the succulent stems. In summary, photosynthesis in stems of submerged plants increased the pO2 in the shoots and roots so that tissues experience diurnal changes in the pO2, but O2 from the H2O column also entered submerged plants.     

36个稀有树种的耐性分析

对湖南南岳树木园内引种的鹅掌楸 [Liriodendron chinense (Hemsl.) Sarg.]、桂南木莲(Manglietia chingii Dandy)等36个稀有树种分别进行了耐干旱、耐水涝和耐阴蔽的研究.结果表明,在全自然光照条件下,各树种的暂时凋萎系数为4.97%～11.05%,凋萎系数为1.85%～5.87%.水淹8～68 d,有31个树种出现叶的第1次凋萎,有22种在水淹30～68 d出现死亡,而长叶竹柏(Podocarpus fleuryi Hickel)等5树种水淹73 d未出现叶凋萎.全阴蔽条件下,阴蔽致死亡树种的时间在17～72 d之间,强阴蔽对树种造成伤害.研究结果可为稀有树种的引种、繁育和管理提供科学依据.     

Growth and Solute Composition in Two Wheat Species Experiencing Combined Influence of Stress Conditions1

The effects of environmental stress combinations on the soluble metabolites were investigated in several cultivars of Triticum aestivum and T. durum. The seedlings grown at optimum (24/16°C), low (5/–5°C) (LT), and high (40/30°C) (HT) day/night temperature conditions were exposed to waterlogging, drought, and salinity (0.7% NaCl, w/w) stresses for six days. Root and shoot fresh weight significantly decreased under waterlogging, drought and salt stresses. Fresh weight was most reduced at severe drought + HT combinations. The lowest relative water content was found under drought stress + HT combination. Soluble carbohydrate (SC) contents increased under LT conditions, but decreased under HT conditions. Under HT + salt combinations, T. aestivum genotypes showed higher SC content thanT. durum genotypes. Proline content significantly increased in the case of water deficit and salt stress. Under drought and salt stresses, T. aestivum genotypes had lower proline contents than T. durum genotypes. These results indicate that biochemical responses to drought, waterlogging, and salt stresses were significantly changed in wheat seedlings under LT and HT conditions.     

New insights into reactive oxygen species and nitric oxide signalling under low oxygen in plants

Plants produce reactive oxygen species (ROS) when exposed to low oxygen (O₂). Much experimental evidence has demonstrated the existence of an oxidative burst when there is an O₂ shortage. This originates at various subcellular sites. The activation of NADPH oxidase(s), in complex with other proteins, is responsible for ROS production at the plasma membrane. Another source of low O₂‐dependent ROS is the mitochondrial electron transport chain, which misfunctions when low O₂ limits its activity. Arabidopsis mutants impaired in proteins playing a role in ROS production display an intolerant phenotype to anoxia and submergence, suggesting a role in acclimation to stress. In rice, the presence of the submergence 1A (SUB1A) gene for submergence tolerance is associated with a higher capacity to scavenge ROS. Additionally, the destabilization of group VII ethylene responsive factors, which are involved in the direct O₂ sensing mechanism, requires nitric oxide (NO). All this evidence suggests the existence of a ROS and NO – low O₂ mechanism interplay which likely includes sensing, anaerobic metabolism and acclimation to stress. In this review, we summarize the most recent findings on this topic, formulating hypotheses on the basis of the latest advances.     

Growth responses to waterlogging and drainage of woody Hakea (Proteaceae) seedlings,originating from contrasting habitats in south-western Australia

We investigated the responses of seven woody Hakea (Proteaceae) species (two populations of each), to two months of waterlogging and subsequent drainage, in a controlled glasshouse experiment. The species originated from contrasting environments (winter-wetland versus non-wetland habitats), and differed in abundance (endangered ironstone species versus common species). Waterlogging arrested growth of the main root system, and stimulated the formation of superficial adventitious roots just below the root/shoot junction in all species. Wetland species produced at least twice the amount of adventitious root dry mass of that of non-wetland species, due to differences in number, length or degree of branching. Their adventitious roots also tended to have higher porosities (7–10% versus 5–6% gas spaces). The relative amount of adventitious roots formed was strongly, positively correlated with the maintenance of shoot growth, and only the non-wetland species showed significant shoot growth reductions (19%) upon waterlogging. Dry mass percentage of stems and leaves, and leaf dry mass per area (LMA) increased considerably during waterlogging in all species (averages of 15, 29 and 27%, respectively), returning to the values of continuously drained control plants after drainage. Similarly, upon drainage, a suppression of shoot growth (average 35%) and a stimulation of root growth (average 50%) restored the root mass ratios to those of control plants. We found a negative correlation between the maintenance of growth during waterlogging versus that after waterlogging, suggesting a trade-off in functioning of the superficial adventitious roots between waterlogged and drained conditions. The rare winter-wet ironstone endemics resembled the common winter-wet species in most of their responses to waterlogging and drainage. Therefore, the results presented here cannot offer an explanation for their different distribution patterns. Our results suggest that non-wetland species may be disadvantaged in a wetland environment, due to their lower capacity to form adventitious roots resulting in stronger growth reductions.     

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

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.     

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

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 O₂ 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 O₂ loss (ROL), taken using root‐sleeving O₂ 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 N₂. This illustrates the importance of O₂ diffusion into the basal regions of roots within an aerobic zone, and the subsequent longitudinal movement of O₂ within the aerenchyma, to supply O₂ to the tip growing in an O₂ deficient zone.     

淹涝胁迫对鹅掌楸属植物叶片部分生理指标的影响

研究了淹涝胁迫对不同种源鹅掌楸[Liriodendron chinense（Hemsl．）Sarg．]和杂种鹅掌楸（L．chinense xL．tulipifera）幼苗部分生理指标的影响。结果表明：在淹水处理24h后,杂种鹅掌楸幼苗叶片的净光合速率变化不明显,鹅掌楸幼苗叶片的净光合速率则有一定程度的下降;在淹水处理48h后,净光合速率均大幅度下降。在淹涝胁迫下及胁迫解除后,幼苗叶片叶绿素含量持续缓慢下降;而脯氨酸含量总体上呈现上升的趋势;与鹅掌楸相比,杂种鹅掌楸幼苗的超氧化物歧化酶（SOD）和过氧化物酶（POD）活性变化不明显,但SOD活性最高,POD活性最低。结果表明,杂种鹅掌楸具有较强的耐淹涝胁迫的能力。     

水淹对互花米草生长及生理的影响

研究了互花米草在不同没顶水淹时间处理下,株高、叶面积等生长指标以及叶片光合速率与色素、脯氨酸、可溶性糖和蛋白质含量等生理指标变化情况。结果表明,随着水淹时间延长,米草株高和叶面积呈下降趋势,叶片光合速率下降;叶片中自由水/束缚水、叶绿素和类胡萝卜素含量、可溶性糖在不同水淹时长处理之间也存在显著差异;但可溶性蛋白质和游离脯氨酸含量在各处理间差异无显著性。