Hormone sensitivity and plant adaptations to flooding

Plant hormones play a key role as mediators between environmental signals and adaptive plant responses. Auxin, ethylene and gibberellins are involved in the initiation of adaptive plant responses such as the development of adventitious roots and stimulated shoot elongation upon flooded conditions. These adaptive plastic responses in plants are frequently linked to changes in the concentrations of the hormones involved, but only rarely to shifts in sensitivity. Examples from ecophysiological research performed with species from the genusRumex demonstrate the importance of the hormone sensitivity concept in plant adaptations to flooding: (a)Rumex species can be grouped into three response categories according to the ethylene sensitivity of the youngest petioles: positive, negative and indifferent; (b) Sub-ambient oxygen concentrations sensitize petioles of wetlandRumex species to ethylene; (c) Enhanced ethylene levels sensitize petioles of wetlandRumex species to gibberellin; (d) Auxin is the primary plant hormone responsible for the initiation of adventitious roots in wetlandRumex species. However, a factor related to waterlogging, possibly ethylene, is required to sensitize the root-shoot junction to endogenous auxin.     

Ethylene accumulation in waterlogged Rumex plants promotes formation of adventitious roots

Accumulation of the gaseous plant hormone ethylene is very importantfor the induction of several responses of plants to flooding.However, little is known about the role of this gas in the formationof flooding-induced adventitious roots. Formation of adventitiousroots in Rumex species is an adaptation of these plants to floodedsoil conditions. The large air-spaces in these roots enablesdiffusion of gases between shoot and roots. Application of ethylene to non-flooded Rumex plants resultedin the formation of adventitious roots. In R. palustris Sm.shoot elongation and epinasty were also observed. The numberof roots in R. thyrsiflorus Fingerh. was much lower than inR. palustris, which corresponds with the inherent differencein root forming capacity between these two species. Ethyleneconcentrations of 1.5–2µI I_{– 1} induced a maximumnumber of roots in both species. Quantification of ethylene escaping from root systems of Rumexplants that were de-submerged after a 24 h submergence periodshowed that average ethylene concentrations in submerged rootsreached 1.8 and 9.1 µl I_–1 in R. palustris and R.thyrsiflorus, respectively. Inhibition of ethylene productionin R. palustris by L--(2-aminoethoxyvinyl)-glycine (AVG) or-aminobutyric acid (AIB) decreased the number of adventitiousroots induced by flooding, indicating that high ethylene concentrationsmay be a prerequisite for the flooding-induced formation ofadventitious roots in Rumex species. Key words: Adventitious roots, epinasty, ethylene, flooding, Rumex, shoot elongation     

Growth and Survival Responses of Rumex Species to Flooded and Submerged Conditions: The Importance of Shoot Elongation, Underwater Photosynthesis and Reserve Carbohydrates

Plants of Rumex thyrsiflorus Fingerh., R. crispus L. and R.maritimus L., which are zoned along a gradient of elevationin a river foreland ecosystem, and differ in their flood-tolerance,were subjected to different flooding levels. Under conditionsof soil flooding, the growth rates of the flood-tolerant R.crispus and R. maritimus were as high as under drained conditions,but that of the flood-intolerant R. thyrsiflorus was halved.Upon submergence, the low elevation species R. maritimus showedrapid shoot elongation; when elongation resulted in a protrusionof leaves above the water surface, the plants survived. Alternatively,underwater photosynthesis also led to a 100% survival of submergedR. maritimus plants, provided that enough inorganic carbon wasmade available in the water. This could be attributed in partto the use of photosynthetically-derived oxygen for root respiration;in a hydroculture experiment, with 5.0 mM CO₂ in the water inthe shoot environment, photosynthetically-derived oxygen contributedmore than 50% to root oxygen consumption at low oxygen concentrationsin the root environment. The intermediately elevated species R. crispus appeared to bemuch more tolerant towards conditions of prolonged total submergence:older plants survived eight weeks submergence in the dark. Thisresponse was explicable in terms of a dormancy-strategy, whichis characterized by a slow consumption of carbohydrates storedin the tap-root. The differential responses of R. maritimusand R. crispus to total submergence reveal the limitations offlood-tolerance and reflect the different life-histories ofthe species. Key words: Photosynthesis, Rumex, submergence, carbohydrates, growth rate, shoot elongation     

An amalgamation between hormone physiology and plant ecology: A review on flooding resistance and ethylene

Both distribution of terrestrial plants and species composition in flood plain communities are strongly influenced by flooding (waterlogging, partial submergence, or submergence). The interaction between a plant's flooding resistance and the seasonal timing, duration, depth, or frequency of flooding often determines plant distribution in flood plains. Flooding may be accompanied by marked physical changes in light, carbon availability, diffusion rate of gases, and density of the environment. Various physiological processes may be affected by these flooding-induced physical changes, including aerobic respiration, photosynthesis, and processes in which light acts as a source of information (e.g., phytochrome photoequilibrium). Certain plant species acclimatize and adapt to these physical changes to relieve the constraints imposed by the flooded environment. Underwater photosynthesis, enhanced shoot elongation, adventitious roots, and aerenchyma formation are typical adaptive responses which are believed to improve the oxygen status of submerged plants. Ethylene and other plant hormones play a central role in the initiation and regulation of most of these adaptive responses, which permit escape from anaerobiosis. Mechanisms of direct tolerance of anaerobic conditions, such as a vigorous fermentative respiratory pathway, are of particular importance when the plant is very deeply submerged, or during the night and when the water is sufficiently turbid to exclude light.Studies on the cosmopolitan genus Rumex, distributed in a flooding gradient on river flood plains, have integrated plant hormone physiology with plant ecology. Rumex species showed a high degree of interspecific variation in ethylene production rates, endogenous ethylene concentrations, ethylene sensitivity, and ethylene-mediated growth responses. The field distribution of Rumex species in flooding gradients is explained in terms of a balance between endogenous ethylene concentrations and sensitivity towards this growth regulator (ethylene economy). Much data has been gathered using a recently developed laser-driven photoacoustic detection technique capable of detecting six parts of ethylene in 10¹² parts air flowing continuously over the plant.     

Life cycle stage and water depth affect flooding-induced adventitious root formation in the terrestrial species Solanum dulcamara

Background and Aims Flooding can occur at any stage of the life cycle of a plant, but often adaptive responses of plants are only studied at a single developmental stage. It may be anticipated that juvenile plants may respond differently from mature plants, as the amount of stored resources may differ and morphological changes can be constrained. Moreover, different water depths may require different strategies to cope with the flooding stress, the expression of which may also depend on developmental stage. This study investigated whether flooding-induced adventitious root formation and plant growth were affected by flooding depth in Solanum dulcamara plants at different developmental stages.Methods Juvenile plants without pre-formed adventitious root primordia and mature plants with primordia were subjected to shallow flooding or deep flooding for 5 weeks. Plant growth and the timing of adventitious root formation were monitored during the flooding treatments.Key Results Adventitious root formation in response to shallow flooding was significantly constrained in juvenile S. dulcamara plants compared with mature plants, and was delayed by deep flooding compared with shallow flooding. Complete submergence suppressed adventitious root formation until up to 2 weeks after shoots restored contact with the atmosphere. Independent of developmental stage, a strong positive correlation was found between adventitious root formation and total biomass accumulation during shallow flooding.Conclusions The potential to deploy an escape strategy (i.e. adventitious root formation) may change throughout a plant’s life cycle, and is largely dependent on flooding depth. Adaptive responses at a given stage of the life cycle thus do not necessarily predict how the plant responds to flooding in another growth stage. As variation in adventitious root formation also correlates with finally attained biomass, this variation may form the basis for variation in resistance to shallow flooding among plants.     

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

Anatomical changes in roots of wheat seedlings (Triticum aestivumL. cv. Hatri) following oxygen deficiency in the rooting mediumwere investigated. The response of the plant to stress was testedat a very early developmental stage when the first adventitiousroots had just emerged. In order to analyze the adaptation ofdifferent roots, respiration rates of the roots 1–3 and4–n were compared with the respiration rates of the totalroot system. Oxygen deficiency was induced either by flushingnutrient solution with nitrogen or flooding of sand. In contrast to plants grown in well aerated media, both stressvariants led to a significant increase of the intercellularspace of the root cortex in seminal and first adventitious roots.Radial cell enlargement of cortical cells near the root tip,cell wall thickenings in flooded sand cultures and an increasein phloroglucinol-stainable substances were found to be furtherindicators of low oxygen supply. The roots 4–n which were promoted in growth under hypoxiashowed higher respiration rates; hence the total root respirationwas not restricted. Triticum aestivum L. cv. Hatri, wheat, roots, anatomy, anaerobiosis, stress, root respiration, intercellular space     

中华蚊母树在干旱-水淹交叉胁迫下形态和活性氧代谢的适应机制

为阐明中华蚊母树(Distylium chinense)在消落带干旱-水淹交叉胁迫下的形态和活性氧(ROS)代谢适应机制,通过控制实验模拟了三峡水库消落带的水文节律,研究了干旱-水淹交叉胁迫及恢复过程施加不同外源物质对中华蚊母树形态学和ROS清除的变化。结果表明：(1)前期干旱胁迫增强了中华蚊母树对后期水淹胁迫的适应,主要表现在叶片脱落、大量不定根的形成及茎基部膨大等形态学的变化;(2)干旱或水淹单一胁迫下,中华蚊母树·OH、■等ROS水平明显高于对照,表现出氧化应激反应,其超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)等抗氧化系统酶活性及脯氨酸(Pro)等抗氧化系统小分子含量也均显著高于对照,表现出一定的抗氧化防御作用机制,且在复合胁迫下,SOD、CAT、APX酶活性及Pro含量显著高于单一胁迫;(3)恢复阶段,相关性分析表明,中华蚊母树清除ROS(·OH、■的酶促(SOD、CAT、APX)及非酶促(Pro)系统具有一定的协同性。同时,恢复阶段施加脱落酸(ABA),内源Pro显著高于正常水平;施加Pro, SOD、CAT等抗氧化酶活性显著高于对照;施加可...     

Regulatory role of auxin in adventitious root formation in two species of Rumex,differing in their sensitivity to waterlogging

Adventitious rooting in Rumex plants, in which the root systems were in hypoxic conditions, differed considerably between two species. R. palustris, a species from frequently flooded river forelands, developed a large number of adventitious roots during hypoxia, whereas adventitious root formation was poor in R. thyrsiflorus, a species from seldom flooded dykes and river dunes. Adventitious rooting could also be evoked in aerated plants of both species by application of auxin (1-naphthaleneacetic acid or indoleacetic acid) to the leaves. The response to auxin was dose-dependent, but even high auxin doses could not stimulate R. thyrsiflorus to produce as many adventitious roots as R. palustris. Consequently, the difference between the species in the amount of adventitious root formation was probably genetically determined, and not a result of a different response to auxin. A prerequisite for hypoxia-induced adventitious root formation is the basipetal transport of auxin within the shoot, as specific inhibition of this transport by N-1-naphthylphthalamic acid severely decreased the number of roots in hypoxia-treated plants. It is suggested that hypoxia of the root system causes stagnation of auxin transport in the root system. This can lead to an accumulation of auxin at the base of the shoot rosette, resulting in adventitious root formation.     

Ethylene Production and Activation of Hydrolytic Enzymes during Acclimation of Maize Seedlings to Partial Flooding

The effects of partial flooding on the partial pressure of oxygen and carbon dioxide in water around the roots, ethylene production by intact maize (Zea mays L.) seedlings, the activities of hydrolytic enzymes (pectinase, xylanase, and cellulase) in adventitious roots, and the growth of adventitious and main roots were studied. Aggravated hypoxia resulted in the accelerated ethylene production and the activation of enzymes destroying cell walls in the adventitious roots; as a result, the latter changed their growth pattern. The conclusion is that the interrelated responses are adaptive ones, and the adventitious roots play a key role in plant adaptation.     

Stomatal Responses of Woody Seedlings to Flooding in Relation to Nutrient Status in Leaves

Ten-week-old woody seedlings of Gmelina arborea Roxb., Tectonagrandis L. (De Wild. & Th. Dur.) Merrill, and Nauclea diderrichiif., were subjected to 10 weeks flooding and a 5-week-long post-floodingperiod. Flooding induced the development of hypertrophied lenticels,adventitious roots, and root aerenchyma in Gmelina and Tectona,while it did not in Nauclea. All three species responded toflooding by first closing their stomata and reopening at differenttimes during the flooding period. The time and extent of reopeningwere earliest and greatest in Gmelina, followed by Tectona,and latest and smallest in Nauclea. Differences in stomatalconductance at the end of the flooding period paralleled theconcentrations of ethanol in roots and inorganic nutrients inleaves. After flooding was discontinued, stomatal conductanceincreased in Gmelina, but did not in the other species. Leafwater potentials and transpiration rates of flooded Gmelinaand Tectona plants were higher at week 8 than at week 2 whilethe reverse was the case in Nauclea. It is argued that whilepersistently small stomatal apertures during the post-floodingperiod in Nauclea may be a strategy to slow down the movementof accumulated toxic substances from the soil, the higher leafwater potential and transpiration in Gmelina and Tectona atweek 8 than at week 2 may represent recovery of shoot waterrelations resulting from adventitious roots development. Key words: Flooding, stomatal conductance, transpiration, adventitious roots, ethanol     

An Ethylene-Mediated Increase in Sensitivity to Auxin Induces Adventitious Root Formation in Flooded Rumex palustris Sm

The hormonal regulation of adventitious root formation induced by flooding of the root system was investigated in the wetland species Rumex palustris Sm. Adventitious root development at the base of the shoot is an important adaptation to flooded conditions and takes place soon after the onset of flooding. Decreases in either endogenous auxin or ethylene concentrations induced by application of inhibitors of either auxin transport or ethylene biosynthesis reduced the number of adventitious roots formed by flooded plants, suggesting an involvement of these hormones in the rooting process. The rooting response during flooding was preceded by increased endogenous ethylene concentrations in the root system. The endogenous auxin concentration did not change during flooding-induced rooting, but a continuous basipetal transport of auxin from the shoot to the rooting zone appeared to be essential in maintaining stable auxin concentrations. These results suggest that the higher ethylene concentration in soil-flooded plants increases the sensitivity of the root-forming tissues to endogenous indoleacetic acid, thus initiating the formation of adventitious roots.     

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

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 cm³ 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.     

水体溶氧影响陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)对完全水淹的耐受力

溶氧是水环境中一个重要的环境因子,为了探讨水中的溶氧含量水平是否会对陆生植物的耐淹能力造成影响,研究了陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)在遭受不同溶氧含量水体完全淹没后的生长表现、存活情况和非结构碳水化合物的变化。实验结果表明:(1)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的存活。受高溶氧水体完全水淹的喜旱莲子草和牛鞭草主茎的完好程度和存活叶的数量均显著高于遭受低溶氧水体完全水淹的喜旱莲子草和牛鞭草,喜旱莲子草和牛鞭草在高溶氧水体完全水淹后的生物量比低溶氧水体完全水淹后要高;(2)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的生长,受高溶氧水体完全水淹的喜旱莲子草主茎伸长生长和不定根生长显著强于受低溶氧水体完全水淹的喜旱莲子草,在不定根的生长上牛鞭草也具有同样的表现。(3)高溶氧水环境有利于减小被完全淹没的喜旱莲子草和牛鞭草的碳水化合物消耗,两种植物在受高溶氧完全水淹后体内具有的非结构性碳水化合物含量均比受低溶氧完全水淹后高。(4)喜旱莲子草比牛鞭草能更好地耐受完全水淹,当处于低溶氧完全水淹时表现得更为明显,本研究表明入侵物种喜旱莲子草比本地物种牛鞭草具有更强的环境适应能力和水淹耐受能力。     

G proteins as regulators in ethylene-mediated hypoxia signaling

Waterlogging or flooding are frequently or constitutively encountered by many plant species. The resulting reduction in endogenous O₂ concentration poses a severe threat. Numerous adaptations at the anatomical, morphological and metabolic level help plants to either escape low oxygen conditions or to endure them. Formation of aerenchyma or rapid shoot elongation are escape responses, as is the formation of adventitious roots. The metabolic shift from aerobic respiration to anaerobic fermentation contributes to a basal energy supply at low oxygen conditions. Ethylene plays a central role in hypoxic stress signaling, and G proteins have been recognized as crucial signal transducers in various hypoxic signaling pathways. The programmed death of parenchyma cells that results in hypoxia-induced aerenchyma formation is an ethylene response. In maize, aerenchyma are induced in the absence of ethylene when G proteins are constitutively activated. Similarly, ethylene induced death of epidermal cells that cover adventitious roots at the stem node of rice is strictly dependent on heterotrimeric G protein activity. Knock down of the unique Gα gene RGA1 in rice prevents epidermal cell death. Finally, in Arabidopsis, induction of alcohol dehydrogenase with resulting increased plant survival relies on the balanced activities of a small Rop G protein and its deactivating protein RopGAP4. Identifying the general mechanisms of G protein signaling in hypoxia adaptation of plants is one of the tasks ahead.Key words: submergence, hypoxia, ethylene, G protein, reactive oxygen species, H₂O₂     

The beneficial effect of reduced elongation growth on submergence tolerance of rice

Adverse effects of elongation growth on tolerance to completesubmergence for up to 14 d were evaluated in rice seedlingsof cultivars which differed in submergence tolerance. Thereis a good negative correlation between per cent survival andelongation growth of genotypes during complete submergence (r= – 0.81). When elongation growth underwater is minimizedby application of a gibberellin biosynthesis inhibitor, percent survival increases by as much as 50 times for one cultivar.These effects are likely related to elongation growth since(i) addition of gibberellin had the opposite effect by reducingsurvival, and (ii) when the elongation inhibitor and gibberellinwere added together, there was no effect on elongation growthand the per cent survival did not change. A GA-deficient mutantof rice which had little elongation ability during submergenceshowed a high level of submergence tolerance when plants weresubmerged at equal initial dry weights and carbohydrate levelsrelative to a submergence-tolerant cultivar. These results areconsistent with the hypothesis that elongation growth competeswith maintenance processes for energy and hence reduces survivalduring submergence. The impact of these findings is that inenvironments where elongation ability is not required, thereis a potential to increase submergence tolerance of agriculturallyimportant cultivars by selecting for least elongation, at leastduring periods of complete submergence. Furthermore, this trade-offbetween stimulated elongation growth and submergence tolerancewill have important ecological consequences for the distributionof plant species in different flood-prone environments. Key words: Gibberellin, growth, Oryza sativa, rice, submergence     

Ethylene enhances gibberellin levels and petiole sensitivity in flooding-tolerant Rumex palustris but not in flooding-intolerant R. acetosa

The role of gibberellin (GA) and ethylene in submergence-induced petiole elongation was studied in two species of the genus Rumex. Analysis of endogenous GAs in the flooding-tolerant Rumex palustris Sm. and the intolerant Rumex acetosa L. by gas chromatography-mass spectrometry showed for both species the presence of GA₁, GA₄, GA₉, GA₁₉, GA₂₀ and GA₅₃. Gas chromatography-mass spectrometry analysis of R. palustris petiole tissue of submerged plants showed an increase in levels of 13-OH GAs, especially GA₁, compared with drained plants. This effect could be mimicked by application of 5 μL L⁻¹ ethylene. In R. acetosa, no differences between levels of GAs in drained or submerged plants were found. In R. palustris, both submergence and ethylene treatment sensitized petioles to exogenous gibberellic acid (GA₃). In R. acetosa the effect was opposite, i.e. submergence and ethylene de-sensitized petioles to GA₃. Our results demonstrate the dual effect of ethylene in the submergence response related to flooding tolerance, i.e. in the flooding-tolerant R. palustris ethylene causes an increased concentration of and sensitivity to GA with respect to petiole elongation while in the intolerant R. acetosa ethylene reduces growth independent of GAs. Received: 5 November 1996 / Accepted: 8 February 1997     

Nitrification and nitrate reductase activity along a secondary successional gradient

The control of nitrification was studied in a secondary successional gradient on Nantucket Island, MA. It was hypothesized that 1) variability in nitrification along the gradient is controlled by litter primary and secondary chemistry, and 2) differences in nitrate availability along the gradient are reflected in potential nitrate assimilation rates in plant tissue. Nitrification varied significantly (p<0.05) by successional stage in all study sites, generally increasing with successional age. The ratio of nitrification to total N mineralization did not vary significantly between successional stages, suggesting substrate limitation of nitrification. Litter terpenoid resin concentration was a significant predictor (p<0.05) of nitrification rate, but soil %C, %N, and water content also contributed significantly to a stepwise regression model predicting nitrification. Nitrate reductase activity (NRA), an index of potential nitrate assimilation, was measured in an assay species (Schizachyrium scoparium). Although there was no significant correlation with nitrification, NRA was significantly (p<0.05) negatively correlated with soil ammonium concentration along the successional gradient at one site, suggesting that plants preferentially utilized ammonium in this system.     

桑苗非结构性碳水化合物和生长激素对水淹胁迫的响应

桑(Morus alba)具有较强的耐水淹特性, 为了探究水淹胁迫对其非结构性碳水化合物和生长激素的影响, 揭示变化规律, 该研究采取室内模拟水淹实验, 以三年生盆栽桑苗作为研究对象, 设置对照组(CK)、根淹组(GY)、浅淹组(QY)、深淹组(SY)等4个不同水淹胁迫的处理, 定期观测并记录桑苗叶片非结构性碳水化合物(可溶性糖和淀粉)含量、内源生长激素(乙烯、脱落酸、赤霉素)含量变化情况。研究结果表明: (1)水淹胁迫会促进桑苗叶片内的生化反应, 造成叶片可溶性糖含量增加。水淹75天, GY、QY、SY桑苗叶片可溶性糖含量较水淹前分别增加182.18%、170.21%和94.16%, 差异显著, 且显著高于CK。水淹胁迫下桑苗叶片淀粉含量在水淹0-50天无显著变化, 水淹75天, GY、QY、SY桑苗叶片淀粉含量较水淹50天分别增加290.84%、244.65%和130.04%, 差异显著, 且显著高于CK。(2)水淹胁迫下桑苗叶片乙烯和赤霉素含量均显著增加, 水淹75天, GY和SY桑苗乙烯含量较水淹前分别增加62.80%和26.78%, 差异显著; GY、QY和SY桑苗赤霉素含量分别增加27.48%、18.02%和25.04%, 差异显著。随着水淹时间增加, GY和SY桑苗乙烯和赤霉素含量总体均呈增加趋势, QY桑苗乙烯和赤霉素含量先增后减, 但仍高于水淹前。水淹胁迫下水淹各组桑苗叶片脱落酸含量随着水淹深度的增加而增加, 水淹75天, QY和SY桑苗叶片脱落酸含量较水淹前分别增加19.20%和36.16%, 差异显著; GY桑苗脱落酸含量无显著变化。上述研究结果表明桑苗可通过调整体内非结构性碳水化合物(可溶性糖和淀粉)的含量和分配, 同时通过积累乙烯、赤霉素、脱落酸等内源激素以适应水淹环境, 具有较强耐淹能力。     

The Effects of Rootzone Salinity and Hypoxia on Shoot and Root Growth in Trifolium Species

The effect of concurrent salinity (0-60 mM NaCl) and rootzonehypoxia (flooding for up to 15 d) on shoot and root growth andshoot ion concentrations of six species of Trifolium (T. subterraneumL., T. fragiferum L., T. michelianum Savi., T. isthmocarpumBot., T. purpureum Lois., and T. repens L.), was studied intwo greenhouse experiments. There was a significant salinityx flooding effect for shoot yield but no significant salinityx flooding x species interaction although individual speciesdiffered significantly (P < 0·001) in their growthresponse to the saline or flooded conditions separately. Concentrationsof Na and Cl in the shoots of all species increased with increasingperiods of saline flooding and there was a significant salinityx flooding interaction. Sodium and Cl concentrations were significantlyhigher (P < 0·001) in T. purpureum, the species inwhich shoot growth was most depressed by saline flooding, thanother species. In T. michelianum, T. fragiferum and T. repens,fresh and dry weight of roots increased with flooding underboth saline and non-saline conditions while in T. subterraneumroot growth decreased. A significant proportion of the increasedroot growth in the first three species occurred as new adventitiousroots. These roots had higher percentages of internal gas spaceswithin the root tissue even in the presence of salinity comparedwith roots from non-flooded conditions. There were also significantlymore gas spaces in the total root tissue in T. fragiferum andT. repens under saline-flooding than in roots of T. subterraneum.Electron micrographs of the root cross sections illustratedthe presence of these gas spaces or aerenchyma. Trifolium fragiferum, T. repens and T. michelianum are morelikely to be suited to growth in soils prone to high salinityand to flooding than are T. subterraneum, T. purpureum and T.ishmocarpum.Copyright 1993, 1999 Academic Press Trifolium subterraneum, Trifolium fragiferum, Trifolium michelianum, Trifolium isthmocarpum, Trifolium purpureum, Trifolium repens, salinity, flooding, hypoxia, adventitious roots, aerenchyma, subterranean clover, white clover, strawberry clover, purple clover, balansa clover