Wheat genotypes show contrasting abilities to recover from anoxia in spite of similar anoxic carbohydrate metabolism

Physiological and metabolic responses to anoxia and reaeration were compared for 4–7-day-old seedlings of 11 genotypes of wheat (Triticum aestivum) with reputed differences in waterlogging tolerance. Genotypes differed in seminal root elongation, and recovery of root tissue K⁺ concentration, during reaeration following 72 h anoxia. Post-anoxic recovery ranged from complete (100% retention of seminal root elongation potential) to almost nil (death of all seminal root apices and inability to recover K⁺ concentration). The anoxia tolerance ranking of the genotypes based on these parameters corresponded with that of their reputed waterlogging tolerance, but with some exceptions. However, the differences in anoxia tolerance of the seedlings could not be explained by differences in capacity for ethanol production. A decreased ability to utilise seed starch reserves under anoxia, due to inadequate levels of -amylase activity at the time anoxia was imposed, was apparent in all genotypes.     

Root Respiration and Carbohydrate Status of Two Wheat Genotypes in Response to Hypoxia

To investigate root respiration and carbohydrate status in relationto waterlogging or hypoxia tolerance, root respiration rateand concentrations of soluble sugars in leaves and roots weredetermined for two wheat (Triticum aestivum L.) genotypes differingin waterlogging-tolerance under hypoxia (5% O₂) and subsequentresumption of full aeration. Root and shoot growth were reducedby hypoxia to a larger extent for waterlogging-sensitive Coker9835. Root respiration or oxygen consumption rate declined withhypoxia, but recovered after 7 d of resumption of aeration.Respiration rate was greater for sensitive Coker 9835 than fortolerant Jackson within 8 d after hypoxia. The concentrationsof sucrose, glucose and fructose decreased in leaves for bothgenotypes under hypoxia. The concentration of these sugars inroots, however, increased under hypoxia, to a greater degreefor Jackson. An increase in the ratio of root sugar concentrationto shoot sugar concentration was found for Jackson under hypoxicconditions, suggesting that a large amount of carbohydrate waspartitioned to roots under hypoxia. The results indicated thatroot carbohydrate supply was not a limiting factor for rootgrowth and respiration under hypoxia. Plant tolerance to waterloggingof hypoxia appeared to be associated with low root respirationor oxygen consumption rate and high sugar accumulation underhypoxic conditions.Copyright 1995, 1999 Academic Press Oxygen consumption rate, sugar accumulation, Triticum aestivum L., waterlogging tolerance     

The impact of limited soil moisture and waterlogging stress conditions on morphological and anatomical root traits in maize (Zea mays L.) hybrids of different drought tolerance

Effects of soil drought or waterlogging on the morphological traits of the root system and internal root anatomy were studied in maize hybrids of different drought tolerance. The investigations comprised quantitative and qualitative analyses of a developed plant root system through determining the number, length and dry matter of the particular components of the root system and some traits of the anatomical structure of the seminal root. Obtained results have demonstrated a relatively broad variation in the habit of the root system. This mainly refers, to the number, length and dry matter of lateral roots, developed by seminal root, seminal adventitious and nodal roots as well as to some anatomical properties of the stele, cortex and metaxylem elements. Plants grown under waterlogging or drought conditions showed a smaller number and less dry matter of lateral branching than plants grown in control conditions. The harmful effect of waterlogging conditions on the growth of roots was greater when compared with that of plants exposed to drought. In the measurements of the root morphological traits, the effect of soil drought on the internal root anatomical characteristic was weaker than the effect of soil waterlogging. The observed effects of both treatments were more distinct in a drought sensitive hybrid Pioneer D than in drought resistant Pioneer C one. The drought resistant hybrid Pioneer C distinguished by a more extensive rooting and by smaller alterations in the root morphology caused by the stress conditions than drought sensitive hybrid Pioneer D one. Also the differences between the resistant and the sensitive maize hybrids were apparent for examined root anatomical traits. Results confirm that the hybrid Pioneer D of a high drought susceptibility was found to be also more sensitive to periodieal soil water excess. A more efficient water use and a lower shoot to root (S:R) ratio were found to be major reasons for a higher stress resistance of the hybrid Pioneer C. The reasons for a different response of the examined hybrids to the conditions of drought or waterlogging may be a more economical water balance and more favourable relations between the shoot and root dimensions in the drought resistant genotype. The observed modifications of the internal root structure caused by water deficit in plant tissues may partly influence on water conductivity and transport within roots. The results suggest that the morphological and anatomical traits of the maize root system may be used in practice as direct or indirect selection criteria in maize breeding.     

Morphological Changes in Wheat Seedlings (Triticum aestivum L.) following Root Anaerobiosis and Partial Pruning of the Root System

Morphological changes of roots and shoots following oxygen deficiencyin the root medium and after partial pruning of the root systemwere analyzed to obtain easily measurable parameters of theadaptive capacity of the root system against stress. Wheat seedlings(Triticum aestivum L. cv. Hatri) were cultivated on nutrientsolution which was either aerated or flushed with nitrogen,or were cultivated on flooded sand. On the third day after grainswelling in two pruning variants, roots 1–3 or 4–8were excised. Root anaerobiosis retarded longitudinal growth and biomass accumulationof the shoot and the seminal roots, and stimulated the developmentof adventitious roots. Partial removal caused a general compensativegrowth of the remaining roots under aerobic conditions. Root pruning plus anaerobiosis exceeded the compensatory capacityof the seedlings and thus caused a strong delay of elongationand biomass accumulation of both roots and shoots, includingdecrease of the root/shoot ratio. Roots became independent ofendosperm reserves on the seventh day under aerobic conditionsthough caryopses were not completely exhausted at this time.Additionally, oxygen deficiency delayed the reserve exhaustionprocess. Triticum aestivum L. cv Hatri, wheat, roots, growth analysis, morphology, anaerobiosis, strees, root pruning, compensatory capacity, caryopsis     

Waterlogging tolerance in the tribe Triticeae: the adventitious roots of Critesion marinum have a relatively high porosity and a barrier to radial oxygen loss

Nine species from the tribe Triticeae – three crop, three pasture and three ‘wild’ wetland species – were evaluated for tolerance to growth in stagnant deoxygenated nutrient solution and also for traits that enhance longitudinal O₂ movement within the roots. Critesion marinum (syn. Hordeum marinum) was the only species evaluated that had a strong barrier to radial O₂ loss (ROL) in the basal regions of its adventitious roots. Barriers to ROL have previously been documented in roots of several wetland species, although not in any close relatives of dryland crop species. Moreover, the porosity in adventitious roots of C. marinum was relatively high: 14% and 25% in plants grown in aerated and stagnant solutions, respectively. The porosity of C. marinum roots in the aerated solution was 1·8–5·4‐fold greater, and in the stagnant solution 1·2–2·8‐fold greater, than in the eight other species when grown under the same conditions. These traits presumably contributed to C. marinum having a 1·4–3 times greater adventitious root length than the other species when grown in deoxygenated stagnant nutrient solution or in waterlogged soil. The length of the adventitious roots and ROL profiles of C. marinum grown in waterlogged soil were comparable to those of the extremely waterlogging‐tolerant species Echinochloa crus‐galli L. (P. Beauv.). The superior tolerance of C. marinum, as compared to Hordeum vulgare (the closest cultivated relative), was confirmed in pots of soil waterlogged for 21 d; H. vulgare suffered severe reductions in shoot and adventitious root dry mass (81% and 67%, respectively), whereas C. marinum shoot mass was only reduced by 38% and adventitious root mass was not affected.     

Morpho-anatomical and physiological responses to waterlogging stress in different barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.) genotypes

Waterlogging is one of the major stresses limiting crop production worldwide. The understanding of the mechanisms of plant adaptations to waterlogging stress helps improve plant tolerance to stress. In this study, physiological responses and morpho-anatomical adaptations of seven different barley genotypes were investigated under waterlogging stress. The results showed that the waterlogging-tolerant varieties (TX9425, Yerong, TF58) showed less reduction in plant height, SPAD (soil–plant analyses development analyses) value, tillers, shoot and root biomasses than did the waterlogging-sensitive varieties (Franklin, Naso Nijo, TF57). Under waterlogging stress condition, the tolerant genotypes also showed a much larger number of adventitious roots than did the sensitive genotypes. More intercellular spaces and better integrated chloroplast membrane structures were observed in the leaves of the waterlogging-tolerant cultivars, which is likely due to increased ethylene content, decreased ABA content and less accumulation of O₂^.?. The ability to form new adventitious roots and intercellular spaces in shoots can also be used as selection criteria in breeding barley for waterlogging tolerance.     

The influence of soil drought and partial waterlogging on water relations of Gmelina arborea seedlings

Summary Stomatal conductance of unstrossed, soil drought, and previously drought (predrought) Gmelina arborea seedlings increased in the morning and decreased before or immediately after midday. In the unstressed and predrought seedlings, leaf water potential decreased with increases in transpiration. In soil drought seedlings, there was some evidence of decreased hydraulic conductivity from soil to the plant, as indicated by the shape in the slope of the water potential/transpiration relationship. Root growth of drought plants was greater than in their unstressed counterparts at the lowest soil segment of a pot. The partial recovery of predrought seedlings was attributed to this subtantial root growth in the lowest soil segment.In the second experiment, Gmelina arborea seedlings were partially waterlogged, by flooding the polyethylene bag to half its length, for a period of 23 days. Waterlogging induced stomatal closure and reduction in leaf water potential but there was some evidence of tolerance to waterlogging towards the end of treatment. Root growth, shoot and root dry weights were slightly reduced below those of controls. After 9 days of waterlogging, adventitious roots began to form which correlated with depletion of soluble sugars in the shoot but with an increase in the roots.It is suggested that the tolerance of Gmelina plants to either soil drought or waterlogging may partly be due to partitioning of the soluble sugars from shoot to roots for production of roots and formation of adventitious roots respectively which are likely to enhance the flow of water from the soils to the plant. Therefore the plant response is very similar under conditions of increased deficits and surplus of soil water.     

The Development of Waterlogging Damage in Young Wheat Plants in Anaerobic Solution Cultures

Anaerobic (anoxic) solution cultures were used to investigatethe effect of a restricted oxygen supply to roots on the developmentof symptoms of waterlogging damage in young wheat plants, especiallyeffects on growth and nutrient uptake by the shoots. Anaerobicconditions produced by bubbling solutions with oxygen-free nitrogengas caused premature senescence of the lower leaves, slowedshoot fresh weight accumulation, and arrested the growth ofthe seminal roots. However the shoot dry weight initially increasedabove that of the aerobic controls. Nutrient accumulation bythe shoot was severely inhibited by anoxia, the uptake of nitrate,phosphate, and potassium being more affected than that of calciumand magnesium. The calculated concentrations in the xylem streamof all these ions (except nitrate) were equal to, or less than,those in the external solution, suggesting that the slow butcontinuous accumulation of nutrients in the shoot could haveoccurred passively by the mass flow of solution across damagedroots in response to transpiration. Aerenchymatous nodal rootsextended into the anoxic solutions to a maximum length of 12cm but there were few produced, and the size of the root systemremained small and may have limited shoot growth. Inclusionof carbon dioxide (10 kPa partial pressure) in the nitrogengas stream had little additional effect on plants to that causedby anoxia alone. All the responses of wheat to the anaerobic solutions were similarto those observed previously in waterlogged soil, indicatingthat many of the early symptoms of waterlogging damage to wheatcan be caused simply by the direct effects of inadequate oxygensupply to the roots. The results are discussed in relation tocurrent views of the mechanisms contributing to waterloggingdamage to plants.     

Kinetin application to roots and its effect on uptake, translocation and distribution of nitrogen in wheat (Triticum aestivum) grown with a split root system

The root systems of wheat seedlings ( Triticum aestivum L. cv. SUN 9E) were pruned to two seminal roots. One of the roots was supplied with a suboptimal level of NO₃, the other was deprived of N. Different levels of kinetin were supplied to the NO₃-deprived roots. Root respiration and the increment of C and N in the roots were measured to determine the C/N ratio of the phloem sap feeding the NO₃-deprived roots. Thus, it was possible to determine retranslocation of N from the shoots to the roots, as affected by the rate of kinetin application. It was calculated that the C/N ratio of phloem sap feeding roots growing without kinetin was ca 61. Kinetin application increased this ratio to ca 75, partly due to decreased translocation of N from the shoots back to the roots. Kinetin application decreased the proportion of N that was retranslocated to the roots after translocation to the shoots. Kinetin increased the rate of NO₃ uptake per root and the rate of N incorporation in both roots and shoots by ca 60%, but had no effect on shoot dry matter production. In control plants at most 70% of the N incorporated in the NO₃-fed roots could have been imported from the shoots, whilst kinetin application reduced this value to ca 40%. Thus root growth was not fully dependent on a supply of N via the phloem.
It is concluded that cytokinins affect the pattern of N-translocation in wheat plants by increasing incorporation of N in dry matter of the shoot, thus leaving less for export. Cytokinins did not play a major role in the regulation of shoot growth and the shoot to root ratio of the present plants.     

小麦根愈伤组织胚胎发育过程研究

实验通过对６个人工合成小麦品系和对照品种“中国春”种子根愈伤组织分化形成再生植株的过程进行形态和组织切片观察,发现分化初期有２种途径,一种是从愈伤组织先形成不定胚,然后再发育成不定芽和不定根,另一种途径是直接从愈伤组织中分化发育成不定根和不定芽;分化后期不定芽和不定根生长发育有３种类型：一种是不定芽发育先于不定根,一种是不定芽与不定期不定芽和不定根生长发育有种类型：一种是不一定芽发育先于不定根,一     

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

Abstract 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.     

Effects of applying etnylene to the root system of Zea mays on growth and nutrient concentration in relation to flooding tolerance

Previous studies have shown increases in the concentration of ethylene in the soil and roots of plants when the soil is water saturated (flooded). In Zea mays L. this occurs in association with an overall reduction in growth but without extensive foliar senescence and in conjunction with the development of an adventitious root system. We have assessed the possibility that ethylene may be involved in these responses to flooding. Mixtures of the gas in air were therefore supplied to the roots and stem-base of Z. mays growing in nutrient solution.
Seven or 14 d exposure to ethylene (1 or 5 νl 1⁻¹) inhibited seminal root elongation and growth in dry weight and accelerated the emergence of adventitious roots, although their final length and dry weight were depressed. Leaf extension was inhibited by 0.1,1.0 or 5.0 μl 1⁻¹ ethylene around the roots; leaves extending rapidiy at the start of treatment were the most sensitive. Final shoot fresh and dry weights were depressed by the gas but tie shootrroot dry weighl ratio and percentage dry matter were not affected greatly. Leaf chlorosis was not observed but the concentration of phosphorus in the shoots was 26 to 31% below normal.
When aeration of the nutrient solution was stopped, the concentration of dissolved oxygen declined and the concentration of ethylene in the roots increased. Similar changes occur in response to soil flooding. Root and shoot growth was slowed by non-aeration although the shootroot dry weight ratio remained unchanged. The phosphorus concentration of the shoots was depressed but there was little chlorosis or leaf death. The similarity in these respects between the effects of ethylene and non-aeration suggests that in flooded Z. mays , ethylene contributes to their development by accelerating the emergence of adventitioos roots, inhibiting phosphorus accumulation in the shoots and by a non-toxic inhibition of plant growth.     

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.     

Flooding tolerance in halophytes

Flooding is a common environmental variable with salinity. Submerged organs can suffer from O₂ deprivation and the resulting energy deficits can compromise ion transport processes essential for salinity tolerance. Tolerance of soil waterlogging in halophytes, as in glycophytes, is often associated with the production of adventitious roots containing aerenchyma, and the resultant internal O₂ supply. For some species, shallow rooting in aerobic upper soil layers appears to be the key to survival on frequently flooded soils, although little is known of the anoxia tolerance in halophytes. Halophytic species that inhabit waterlogged substrates are able to regulate their shoot ion concentrations in spite of the hypoxic (or anoxic) medium in which they are rooted, this being in stark contrast with most other plants which suffer when salinity and waterlogging occur in combination. Very few studies have addressed the consequences of submergence of the shoots by saline water; these have, however, demonstrated tolerance of temporary submergence in some halophytes.     

Functional linkage between N acquisition strategies and aeration capacities of hydrophytes for efficient oxygen consumption in roots

We evaluated the specific strategies of hydrophytes for root O₂ consumption in relation to N acquisition and investigated whether the strategies varied depending on the aeration capacity. Aeration capacity of roots is an important factor for determining hypoxia tolerance in plants. However, some hydrophytes possessing quite different aeration capacities often co‐occur in wetlands, suggesting that root O₂ consumption also strongly affects hypoxia tolerance. We cultivated Phragmites australis with high aeration capacity and Zizania latifolia with low aeration capacity in hypoxic conditions with NH or NO treatment and compared the growth, N uptake, N assimilation and root respiration between the two species. In Z. latifolia grown with NH treatment, high N uptake activity and restrained root growth led to sufficient N acquisition and decrease in whole‐root respiration rate. These characteristics consequently compensated for the low aeration capacity. In contrast, in P. australis, low N uptake activity was compensated by active root growth, but the whole‐root respiration rate was high. This high root respiration rate was allowed by the high aeration capacity. The O₂ consumption‐related traits of hydrophyte roots were closely correlated with N acquisition strategies, which consequently led to a compensational relationship with the root aeration capacity. It is likely that this functional linkage plays an important role as a core mechanism in the adaptation of plants to hypoxic soils.     

Supply and partitioning of assimilates to roots of Medicago sativa L. and Lotus corniculatus L. under anoxia

Abstract A current explanation of the mechanism of flooding injury to roots suggests that oxygen deficiency depresses the supply of respirable carbohydrates sufficiently to inhibit fermentation. However, even though it has been shown that phloem transport of assimilate is sharply reduced to anaerobic roots, inhibition of assimilate metabolism has also been suggested to be an important factor. This study examines these hypotheses by relating assimilate supply and metabolic activity in anoxic roots of alfalfa (Medicago sativa L.), a flood-intolerant species, and birdsfoot trefoil (Lotus corniculatus L.), a flood-tolerant plant. Roots were made anoxic (severe O₂ deficiency) for 2, 4 or 6 d and shoots were labelled with ¹⁴CO₂. Assimilate transport to the roots and metabolism to structural components were significantly decreased in both species in response to anoxia. Trefoil exhibited significantly greater ¹⁴C incorporation into the residue fraction at 4 d anoxia than did alfalfa, and this was consistent with the greater flooding tolerance of trefoil. When assimilate supply to O₂-deficient roots was decreased by shoot shading, shoot fresh weight was reduced by both anoxia and light treatments. Root-soluble sugars were significantly decreased by shading but were greatly increased in response to anoxia. Root starch concentration also increased under anoxia. Root K⁺ concentration was reduced by anoxia only. The energy status (ATP/ADP) of roots was significantly decreased by shading; however, anoxia reduced the energy status only in unshaded plants. The data indicate that carbohydrate supply to anaerobic roots does not appear to be a limiting factor in the metabolic response of alfalfa roots. Alternatively, metabolism of assimilate in anoxic roots may be an important determinant of survival.     

Genotypic variation in seedling root architectural traits and implications for drought adaptation in wheat (Triticum aestivum L.)

Root system characteristics are of fundamental importance to soil exploration and below-ground resource acquisition. Root architectural traits determine the in situ space-filling properties of a root system or root architecture. The growth angle of root axes is a principal component of root system architecture that has been strongly associated with acquisition efficiency in many crop species. The aims of this study were to examine the extent of genotypic variability for the growth angle and number of seminal roots in 27 current Australian and 3 CIMMYT wheat (Triticum aestivum L.) genotypes, and to quantify using fractal analysis the root system architecture of a subset of wheat genotypes contrasting in drought tolerance and seminal root characteristics. The growth angle and number of seminal roots showed significant genotypic variation among the wheat genotypes with values ranging from 36 to 56 (degrees) and 3 to 5 (plant⁻¹), respectively. Cluster analysis of wheat genotypes based on similarity in their seminal root characteristics resulted in four groups. The group composition reflected to some extent the genetic background and environmental adaptation of genotypes. Wheat cultivars grown widely in the Mediterranean environments of southern and western Australia generally had wider growth angle and lower number of seminal axes. In contrast, cultivars with superior performance on deep clay soils in the northern cropping region, such as SeriM82, Baxter, Babax, and Dharwar Dry exhibited a narrower angle of seminal axes. The wheat genotypes also showed significant variation in fractal dimension (D). The D values calculated for the individual segments of each root system suggested that, compared to the standard cultivar Hartog, the drought-tolerant genotypes adapted to the northern region tended to distribute relatively more roots in the soil volume directly underneath the plant. These findings suggest that wheat root system architecture is closely linked to the angle of seminal root axes at the seedling stage. The implications of genotypic variation in the seminal root characteristics and fractal dimension for specific adaptation to drought environment types are discussed with emphasis on the possible exploitation of root architectural traits in breeding for improved wheat cultivars for water-limited environments.     

MESOCOTYL ROOT FORMATION IN ECHINOCHLOA PHYLLOPOGON (POACEAE) IN RELATION TO ROOT ZONE AERATION

Echinochloa phyllopogon was grown hydroponically under four root zone gassing treatments to determine aeration effects on the growth and development of the plant root system. Although mesocotyl growth and the number of nodal roots were unaffected by the treatments, other aspects of plant growth were altered. Shoot growth was reduced by hypoxic (5 kPa partial pressure O₂ in nitrogen gas) and anoxic conditions (O₂ free nitrogen gas), but not by ethylene (0.1 ppm in air). Seminal root growth was unaffected by hypoxia or ethylene treatments, but was reduced under anoxia. Hypoxic environments stimulated the emergence of roots along the length of the mesocotyl when compared to aerobic controls; anoxic and ethylene treatments had no significant effects. Mesocotyl roots elongated from primordia that were produced de novo in response to the hypoxic treatment. Under hypoxic conditions, aerenchyma was present in the cortex of nodal roots and to a lesser extent in seminal roots, but mesocotyl roots were devoid of aerenchyma under these conditions. The results are compared with the literature concerning flooding and aeration effects on growth and development in other species.     

Aerenchymatous phellem in hypocotyl and roots enables O2 transport in Melilotus siculus

? Aerenchymatous phellem (secondary aerenchyma) has rarely been studied in roots. Its formation and role in internal aeration were evaluated for Melilotus siculus, an annual legume of wet saline land. ? Plants were grown for 21 d in aerated or stagnant (deoxygenated) agar solutions. Root porosity and maximum diameters were measured after 0, 7, 14 and 21 d of treatment. Phellem anatomy was studied and oxygen (O(2)) transport properties examined using methylene blue dye and root-sleeving O(2) electrodes. ? Interconnecting aerenchymatous phellem developed in hypocotyl, tap root and older laterals (but not in aerial shoots), with radial intercellular connections to steles. Porosity of main roots containing phellem was c. 25%; cross-sectional areas of this phellem were threefold greater for stagnant than for aerated treatments. Root radial O(2) loss was significantly reduced by complete hypocotyl submergence; values approached zero after disruption of hypocotyl phellem below the waterline or, after shoot excision, by covering hypocotyl phellem in nontoxic cream. ? Aerenchymatous phellem enables hypocotyl-to-root O(2) transport in M. siculus. Phellem increases radially under stagnant conditions, and will contribute to waterlogging tolerance by enhancing root aeration. It seems likely that with hypocotyl submerged, O(2) will diffuse via surface gas-films and internally from the shoot system.