Formation and function of secondary aerenchyma in hypocotyl, roots and nodules of soybean (Glycine max) under flooded conditions

Flooding is a major problem in many areas of the world and soybean is susceptible to the stress. Understanding the morphological mechanisms of flooding tolerance is important for developing flood-tolerant genotypes. We investigated secondary aerenchyma formation and function in soybean (Glycine max) seedlings grown under flooded conditions. Secondary aerenchyma, a white and spongy tissue, was formed in the hypocotyl, tap root, adventitious roots and root nodules after 3 weeks of flooding. Under irrigated conditions aerenchyma development was either absent or rare and phellem was formed in the hypocotyl, tap root, adventitious roots and root nodules. Secondary meristem partially appeared at the outer parts of the interfascicular cambium and girdled the stele, and then cells differentiated to construct secondary aerenchyma in the flooded hypocotyl. These morphological changes proceeded for 4 days after the initiation of the flooding. After 14 days of treatment, porosity exceeded 30% in flooded hypocotyl with well-developed secondary aerenchyma, while it was below 10% in hypocotyl of irrigated plants that had no aerenchyma. When Vaseline was applied to the hypocotyl of plants from a flooded treatment to prevent the entry of atmospheric oxygen into secondary aerenchyma, plant growth, especially that of roots, was sharply inhibited. Thus secondary aerenchyma might be an adaptive response to flooding.     

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.     

Inhibition by silver ions of gas space (aerenchyma) formation in adventitious roots of Zea mays L. subjected to exogenous ethylene or to oxygen deficiency

We have studied the role of ethylene in accelerating the lytic formation of gas spaces (aerenchyma) in the cortex of adventitious roots of maize (Zea mays L.) growing in poorly aerated conditions. Such roots had previously been shown to contain increased concentrations of ethylene. Ten day-old maize plants bearing seminal roots and one whorl of emerging adventitious roots were grown in nutrient solution bubbled with air, ethylene in air (0.1 to 5.0 l l^–1), or allowed to become oxygen-deficient in nonaerated (but not completely anaerobic) solution. Additions of 0.1 l l^–1 ethylene or more promoted the formation of aerenchyma, with lysis of up to 47% of the cortical cells. The effects of non-aeration were similar to those of exogenous ethylene. When silver ions, an ethylene antagonist, were present at low, non-toxic concentrations (circa 0.6 M), aerenchyma formation was prevented in ethylene treated roots and in those exposed to oxygen deficiency. Silver ions also blocked the inhibiting effect of exogenous ethylene on root extension. By contrast, the suppression of aerenchyma formation by silver ions under oxygendeficient conditions was associated with a retardation of root extension, indicating the importance of aerenchyma for root growth in poorly aerated media. Rates of production of ethylene by excised roots were stimulated by a previous non-aeration treatment. The effectiveness of Ag⁺ in inhibiting equally the action on cortical cells of exogenous ethylene and of non-aeration, supports the view that gas space (aerenchyma) formation in adventitious roots adpted to oxygendeficient environments is mediated by increased concentrations of endogenous ethylene. The possibility that extra ethylene could arise from increased biosynthesis of a precursor in root tissues with a restricted oxygen supply is discussed.     

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.     

洪水条件下湿地植物的生存策略

洪水是自然界存在的一种普遍现象。湿地植物由于所处生境的特殊性,会经常受到周期性或永久性的洪水胁迫。在长期的适应进化过程中,湿地植物形成了一些特殊的生存策略,以适应水文条件的大幅度变化。主要的生存策略如下:1)生活史方面,植物可通过改变生长时间、繁殖方式、种子特征等避免洪水的直接伤害或利用洪水的流动起到传播扩散的作用;2)形态学特征方面,植物可通过调整根系形态、分布等将根系生长到氧气相对充足的土壤表层或形成不定根增强根系通气功能;3)解剖学方面,植物可通过改善组织孔隙度形成通气组织等改善空气传导到根系的"气体通道";4)生理生化方面,植物可通过增加碳水化合物含量以延长生存时间,释放出一些生长激素(乙烯等)以调节植物缺氧条件下的生理活动或形态、解剖方面的变化。在今后的研究中,不定根的形成机理、乙烯在通气组织形成中的作用及其过程、放射氧损失(ROL)的形成机理及其释放速率的调控等一些机理性的工作还需进一步加强。     

Variation for Root Aerenchyma Formation in Flooded and Non-Flooded Maize and Teosinte Seedlings

Morphological and anatomical factors such as aerenchyma formation in roots and the development of adventitious roots are considered to be amongst the most important developmental characteristics affecting flooding tolerance. In this study we investigated the lengths of adventitious roots and their capacity to form aerenchyma in three- and four-week-old seedlings of two maize (Zea mays ssp. mays, Linn.) inbred accessions, B64 and Na4, and one teosinte, Z. nicaraguensis Iltis & Benz (Poaceae), with and without a flooding treatment. Three weeks after sowing and following a seven day flooding treatment, both maize and teosinte seedlings formed aerenchyma in the cortex of the adventitious roots of the first three nodes. The degree of aerenchyma formation in the three genotypes increased with a second week of flooding treatment. In drained soil, the two maize accessions failed to form aerenchyma. In Z. nicaraguensis, aerenchyma developed in roots located at the first two nodes three weeks after sowing. In the fourth week, aerenchyma developed in roots of the third node, with a subsequent increase in aerenchyma in the second node roots. In a second experiment, we investigated the capacity of aerenchyma to develop in drained soil. An additional three teosinte species and 15 maize inbred lines, among them a set of flooding-tolerant maize lines, were evaluated. Evaluations indicate that accessions of Z. luxurians (Durieu & Asch. Bird) and two maize inbreds, B55 and Mo20W, form aerenchyma when not flooded. These materials may be useful genetic resources for the development of flooding-tolerant maize accessions.     

Enhanced formation of aerenchyma and induction of a barrier to radial oxygen loss in adventitious roots of Zea nicaraguensis contribute to its waterlogging tolerance as compared with maize (Zea mays ssp. mays)

Enhancement of oxygen transport from shoot to root tip by the formation of aerenchyma and also a barrier to radial oxygen loss (ROL) in roots is common in waterlogging‐tolerant plants. Zea nicaraguensis (teosinte), a wild relative of maize (Zea mays ssp. mays), grows in waterlogged soils. We investigated the formation of aerenchyma and ROL barrier induction in roots of Z. nicaraguensis, in comparison with roots of maize (inbred line Mi29), in a pot soil system and in hydroponics. Furthermore, depositions of suberin in the exodermis/hypodermis and lignin in the epidermis of adventitious roots of Z. nicaraguensis and maize grown in aerated or stagnant deoxygenated nutrient solution were studied. Growth of maize was more adversely affected by low oxygen in the root zone (waterlogged soil or stagnant deoxygenated nutrient solution) compared with Z. nicaraguensis. In stagnant deoxygenated solution, Z. nicaraguensis was superior to maize in transporting oxygen from shoot base to root tip due to formation of larger aerenchyma and a stronger barrier to ROL in adventitious roots. The relationships between the ROL barrier formation and suberin and lignin depositions in roots are discussed. The ROL barrier, in addition to aerenchyma, would contribute to the waterlogging tolerance of Z. nicaraguensis.     

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.     

Ethylene-induced activation of xylanase in adventitious roots of maize as a response to the stress effect of root submersion]

Submersion of roots of ten-day-old maize (Zea mays L.) seedlings was accompanied by a decrease in pO2 and an increase in pCO2 of the medium adjacent to roots. These changes stimulated ethylene evolution in intact plants. Enhanced biosynthesis of ethylene was accompanied by xylanase activation in adventitious roots. As a result, an enhanced formation of aerenchyma was observed in the cortex of adventitious roots. Therefore, these processes resulted in the development of a ventilation system by which O2 can reach the root system exposed to hypoxia. The volume of aerenchyma was assessed by the volume of gas cavities (porosity). In contrast to the main root, the growth of adventitious roots was not inhibited under these conditions. Enlargement of the stem base and increase in the number of aerenchymatous adventitious roots facilitated the oxygen supply to submerged organs of plants.     

Dynamics of Aerenchyma distribution in the cortex of sulfate-deprived adventitious roots of maize

BACKGROUND AND AIMS: Aerenchyma formation in maize adventitious roots is induced in nutrient solution by the deprivation of sulfate (S) under well-oxygenated conditions. The aim of this research was to examine the extent of aerenchyma formation in the cortex of sulfate-deprived adventitious roots along the root axis, in correlation with the presence of reactive oxygen species (ROS), calcium levels and pH of cortex cells and root lignification. METHODS: The morphometry of the second whorl of adventitious (W2) roots, subject to S-deprivation conditions throughout development, was recorded in terms of root length and lateral root length and distribution. W2 roots divided into sectors according to the mean length of lateral roots, and cross-sections of each were examined for aerenchyma. In-situ detection of alterations in ROS presence, calcium levels and pH were performed by means of fluorescence microscopy using H(2)DCF-DA, fluo-3AM and BCECF, respectively. Lignification was detected using the Wiesner test. KEY RESULTS: S-deprivation reduced shoot growth and enhanced root proliferation. Aerenchyma was found in the cortex of 77 % of the root length, particularly in the region of emerging or developing lateral roots. The basal and apical sectors had no aerenchyma and no aerenchyma connection was found with the shoot. S-deprivation resulted in alterations of ROS, calcium levels and pH in aerenchymatous sectors compared with the basal non-aerenchymatous region. Lignified epidermal layers were located at the basal and the proximal sectors. S-deprivation resulted in shorter lateral roots in the upper sectors and in a limited extension of the lignified layers towards the next lateral root carrying sector. CONCLUSIONS: Lateral root proliferation is accompanied by spatially localized induced cell death in the cortex of developing young maize adventitious roots during S-deprivation.     

Cortical Aerenchyma formation in hypocotyl and adventitious roots of Luffa cylindrica subjected to soil flooding

BACKGROUND AND AIMS: Aerenchyma formation is thought to be one of the important morphological adaptations to hypoxic stress. Although sponge gourd is an annual vegetable upland crop, in response to flooding the hypocotyl and newly formed adventitious roots create aerenchyma that is neither schizogenous nor lysigenous, but is produced by radial elongation of cortical cells. The aim of this study is to characterize the morphological changes in flooded tissues and the pattern of cortical aerenchyma formation, and to analyse the relative amount of aerenchyma formed. METHODS: Plants were harvested at 16 d after the flooding treatment was initiated. The root system was observed, and sections of fresh materials (hypocotyl, tap root and adventitious root) were viewed with a light or fluorescence microscope. Distributions of porosity along adventitious roots were estimated by a pycnometer method. KEY RESULTS: Under flooded conditions, a considerable part of the root system consisted of new adventitious roots which soon emerged and grew quickly over the soil surface. The outer cortical cells of these roots and those of the hypocotyl elongated radially and contributed to the development of large intercellular spaces. The elongated cortical cells of adventitious roots were clearly T-shaped, and occurred regularly in mesh-like lacunate structures. In these positions, slits were formed in the epidermis. In the roots, the enlargement of the gas space system began close to the apex in the cortical cell layers immediately beneath the epidermis. The porosity along these roots was 11-45 %. In non-flooded plants, adventitious roots were not formed and no aerenchyma developed in the hypocotyl or tap root. CONCLUSIONS: Sponge gourd aerenchyma is produced by the unique radial elongation of cells that make the expansigeny. These morphological changes seem to enhance flooding tolerance by promoting tissue gas exchange, and sponge gourd might thereby adapt to flooding stress.     

Ethylene-Induced Activation of Xylanase in Adventitious Roots of Maize as a Response to the Stress Effect of Root Submersion

Submersion of roots of ten-day-old maize (Zea maysL.) seedlings was accompanied by a decrease in pO₂and an increase in pCO₂of the medium adjacent to the roots. These changes stimulated ethylene evolution in intact plants. Enhanced biosynthesis of ethylene was accompanied by xylanase activation in adventitious roots. As a result, an enhanced formation of aerenchyma was observed in the cortex of adventitious roots. Therefore, these processes resulted in the development of a ventilation system by which O₂can reach the root system exposed to hypoxia. The volume of aerenchyma was assessed by the volume of gas cavities (porosity). In contrast to the main root, the growth of adventitious roots was not inhibited under these conditions. Enlargement of the stem base and increase in the number of aerenchymatous adventitious roots facilitated the oxygen supply to the submerged organs of the plants.     

Physiological,morphological, and growth responses ofPlatanus occidentalis seedlings to flooding

Summary Flooding ofPlatanus occidentalis L. seedlings for up to 40 days induced several changes including early stomatal closure, greatly accelerated ethylene production by stems, formation of hypertrophied lenticels and adventitious roots on submerged portions of stems, and marked growth inhibition. Poor adaptation ofPlatanus occidentalis seedlings to soil inundation was shown in stomatal closure during the entire flooding period, inhibition of root elongation and branching, and death of roots. Some adaptation to flooding was indicated by (1) production of hypertrophied lenticels which may assist in exchange of dissolved gases in flood water and in release of toxic compounds, and (2) production of adventitious roots on stems which may increase absorption of water. These adaptations appeared to be associated with greatly stimulated ethylene production in stems of flooded plants. The greater reduction of root growth over shoot growth in flooded seedlings will result in decreased drought tolerance after the flood waters recede. The generally low tolerance to flooding of seedlings of species that are widely rated as highly flood tolerant is emphasized.     

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.     

Arbuscular mycorrhizal colonization and nodulation improve flooding tolerance in <Emphasis Type="Italic">Pterocarpus officinalis</Emphasis> Jacq. seedlings

Pterocarpus officinalis (Jacq.) seedlings inoculated with the arbuscular mycorrhizal fungus, Glomus intraradices, and the strain of Bradyrhizobium sp. (UAG 11A) were grown under stem-flooded or nonflooded conditions for 13 weeks after 4 weeks of nonflooded pretreatment under greenhouse conditions. Flooding of P. officinalis seedlings induced several morphological and physiological adaptive mechanisms, including formation of hypertrophied lenticels and aerenchyma tissue and production of adventitious roots on submerged portions of the stem. Flooding also resulted in an increase in collar diameter and leaf, stem, root, and total dry weights, regardless of inoculation. Under flooding, arbuscular mycorrhizas were well developed on root systems and adventitious roots compared with inoculated root systems under nonflooding condition. Arbuscular mycorrhizas made noteworthy contributions to the flood tolerance of P. officinalis seedlings by improving plant growth and P acquisition in leaves. We report in this study the novel occurrence of nodules connected vascularly to the stem and nodule and arbuscular mycorrhizas on adventitious roots of P. officinalis seedlings. Root nodules appeared more efficient fixing N₂ than stem nodules were. Beneficial effect of nodulation in terms of total dry weight and N acquisition in leaves was particularly noted in seedlings growing under flooding conditions. There was no additive effect of arbuscular mycorrhizas and nodulation on plant growth and nutrition in either flooding treatment. The results suggest that the development of adventitious roots, aerenchyma tissue, and hypertrophied lenticels may play a major role in flooded tolerance of P. officinalis symbiosis by increasing oxygen diffusion to the submerged part of the stem and root zone, and therefore contribute to plant growth and nutrition.     

Stimulation of ethylene production and gas-space (aerenchyma) formation in adventitious roots of Zea mays L. by small partial pressures of oxygen

Adventitious roots of two to four-weekold intact plants of Zea mays L. (cv. LG11) were shorter but less dense after extending into stagnant, non-aerated nutrient solution than into solution continuously aerated with air. Dissolved oxygen in the non-aerated solutions decreased from 21 kPa to 3–9 kPa within 24 h. When oxygen partial pressures similar to those found in non-aerated solutions (3, 5 and 12 kPa) were applied for 7 d to root systems growing in vigorously bubbled solutions, the volume of gas-space in the cortex (aerenchyma) was increased several fold. This stimulation of aerenchyma was associated with faster ethylene production by 45-mm-long apical root segments. When ethylene production by roots exposed to 5 kPa oxygen was inhibited by aminoethoxyvinylglycine (AVG) dissolved in the nutrient solution, aerenchyma formation was also retarded. The effect of AVG was reversible by concomitant applications of 1-aminocyclopropane-1-carboxylic acid, an immediate precursor of ethylene. Addition of silver nitrate, an inhibitor of ethylene action, to the nutrient solution also prevented the development of aerenchyma in roots given 5 kPa oxygen. Treating roots with only 1 kPa oxygen stimulated ethylene production but failed to promote gas-space formation. These severely oxygen-deficient roots seemed insensitive to the ethylene produced since a supplement of exogeneous ethylene that promoted aerenchyma development in nutrient solution aerated with air (21 kPa oxygen) failed to do so in nutrient solution supplied with 1 kPa oxygen. Both ethylene production and aerenchyma formation were almost completely halted when roots were exposed to nutrient solutions devoid of oxygen. Thus both processes require oxygen and are stimulated by oxygen-deficient surroundings in the 3-to 12-kPa range of oxygen partial pressures when compared with rates observed in air (21 kPa oxygen).Abbreviations ACC 1-aminocyclopropane-1-carboxylic acid - AVG aminoethoxyvinylglycine     

Decreased Ethylene Biosynthesis, and Induction of Aerenchyma, by Nitrogen- or Phosphate-Starvation in Adventitious Roots of Zea mays L

Plants of Zea mays L. cv TX5855 were grown in a complete, well oxygenated nutrient solution then subjected to nutrient starvation by omitting either nitrate and ammonium or phosphate from the solution. These treatments induced the formation of aerenchyma close to the apex of the adventitious roots that subsequently emerged from the base of the shoot, a response similar to that shown earlier to be induced by hypoxia. Compared with control plants supplied with all nutrients throughout, N- or P-starvation consistently depressed the rates of ethylene release by excised, 25 mm apical segments of adventitious roots. Some enzymes and substrates of the ethylene biosynthetic pathway were examined. The content of 1-amino cyclopropane-1-carboxylic acid (ACC) paralleled the differences in ethylene production rates, being depressed by N or P deficiency, while malonyl-ACC showed a similar trend. Activity of ACC synthase and of ethylene forming enzyme (g⁻¹ fresh weight) was also greater in control roots than in nutrient starved ones. These results indicate that much of the ethylene biosynthetic pathway is slowed under conditions of N- or P-starvation. Thus, by contrast to the effects of hypoxia, the induction of aerenchyma in roots of Zea mays by nutrient starvation is not related to an enhanced biosynthesis and/or accumulation of ethylene in the root tips.     

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     

Hormones and root-shoot relationships in flooded plants — an analysis of methods and results

Two aspects of root to shoot communication in flooded plants are discussed (i) the formation of porous aerenchyma that enhances the passage of oxygen, and other gases, from shoots to roots and (ii) the movement of the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) from roots to shoots in the transpiration stream, and the effect of this on ethylene production and epinastic curvature in the shoots. For aerenchyma studies a highly sensitive photoacoustic laser detector for ethylene was used to avoid interference associated with other methods of ethylene measurement that require tissue excision. ACC concentrations in xylem sap were measured by physico-chemical means to ensure correct identification and account for processing losses. Solute concentrations, e.g., abscisic acid (ABA), in xylem sap are shown to be distorted by temporary contamination caused by the method used to collect sap. Concentrations of solutes in xylem sap (e.g., ACC) are also altered by changes in sap flow brought about by conventional methods of sap collection or by experimental treatments such as flooding the soil. Ways of for overcoming these problems are described together with a summary of preliminary results.     

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.