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.     

Growth Responses and Adaptations of Fraxinus pennsylvanica Seedlings to Flooding

Flooding induced several physiological and morphological changes in Fraxinus pennsylvanica seedlings, with stomatal closure among the earliest responses. Subsequent changes included: reduction in dry weight increment of roots, stems, and leaves; formation of hypertrophied lenticels and production of adventitious roots on submerged portions of the stem above the soil line; leaf necrosis; and leaf abscission. After 15 days of stomatal closure as a result of flooding, stomata began to reopen progressively until stomatal aperture was similar in flooded and unflooded plants. Adventitious roots began to form at about the time stomatal reopening began. As more adventitious roots formed, elongated, and branched, the stomata opened further. The formation of adventitious roots was an important adaptation for flooding tolerance as shown by the high efficiency of adventitious roots in absorption of water and in high correlation between the production of adventitious roots and stomatal reopening.     

Some physiological and growth responses of kiwi fruit (Actinidia chinensis) to flooding

The effects of long-term flooding on the growth of six-month-old Actinidia chinensis Planch cv. Abbot plants and some effects on stomatal behaviour and leaf water relations were examined under controlled conditions for 28 days. Flooding caused stomatal closure and decreases in transpiration rate, xylem water potential, osmotic potential and turgor potential. Flooding also caused inhibition of the dry weight increase of leaves plus stems and of roots, chlorosis and necrosis of leaves, production of hypertrophied lenticels and the appearance of a small number of adventitious roots on the submerged portions of the stems. Rapid and partial stomatal closure by flooding may not only be due to the passive mechanical response which follows leaf dehydration, since flooded plants showed an increase in xylem water potential and osmotic potential during the first days of the experiment. The marked intolerance of Actinidia chinensis to flooding has been a serious barrier to its culture in poorly drained soils, hence careful irrigation management is required.     

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.     

Erythrina speciosa (Leguminosae-Papilionoideae) under soil water saturation: morphophysiological and growth responses

Background and Aims

Erythrina speciosa is a Neotropical tree that grows mainly in moist habitats. To characterize the physiological, morphological and growth responses to soil water saturation, young plants of E. speciosa were subjected experimentally to soil flooding.

Methods

Flooding was imposed from 2 to 4 cm above the soil surface in water-filled tanks for 60 d. Non-flooded (control) plants were well watered, but never flooded. The net CO₂ exchange (A_CO2), stomatal conductance (g_s) and intercellular CO₂ concentration (C_i) were assessed for 60 d. Soluble sugar and free amino acid concentrations and the proportion of free amino acids were determined at 0, 7, 10, 21, 28 and 45 d of treatments. After 28, 45 and 60 d, dry masses of leaves, stems and roots were determined. Stem and root cross-sections were viewed using light microscopy.

Key Results

The A_CO2 and g_s were severely reduced by flooding treatment, but only for the first 10 d. The soluble sugars and free amino acids increased until the tenth day but decreased subsequently. The content of asparagine in the roots showed a drastic decrease while those of alanine and γ-aminobutyric increased sharply throughout the first 10 d after flooding. From the 20th day on, the flooded plants reached A_CO2 and g_s values similar to those observed for non-flooded plants. These events were coupled with the development of lenticels, adventitious roots and aerenchyma tissue of honeycomb type. Flooding reduced the growth rate and altered carbon allocation. The biomass allocated to the stem was higher and the root mass ratio was lower for flooded plants when compared with non-flooded plants.

Conclusions

Erythrina speciosa showed 100 % survival until the 60th day of flooding and was able to recover its metabolism. The recovery during soil flooding seems to be associated with morphological alterations, such as development of hypertrophic lenticels, adventitious roots and aerenchyma tissue, and with the maintenance of neutral amino acids in roots under long-term exposure to root-zone O₂ deprivation.Key words: Erythrina speciosa, aerenchyma, amino acid content, biomass allocation, photosynthesis, flooding adaptations, stomatal conductance, O₂ deficiency, γ-aminobutyric acid (GABA)     

Importance of adventitious roots to growth of floodedPlatanus occidentalis seedlings

Summary Flooding of soil with standing water for 50 or 110 days drastically reduced growth of 178-day-oldPlatanus occidentalis seedlings, with growth inhibited more as the duration of flooding was increased. Flooding reduced the rate of height and diameter growth, leaf initiation and expansion, and dry weight increment and relative growth rates of leaves, stems, and roots. Flooding also induced leaf epinasty, leaf necrosis, and formation of hypertrophied lenticels and many adventitious roots on submerged portions of stems. Severing of adventitious roots after 50 and 95 days from the submerged portions of stems of continuously flooded seedlings reduced several growth parameters including height and stem diameter growth and relative growth rates of leaves and roots. Evidence for the physiological importance of flood induced adventitious roots is discussed.Research supported by College of Agricultural and Life Sciences, University of Wisconsin, Madison and by Yamagata University, Tsuruoka, Japan. The technical assistance of John Shanklin is appreciated.     

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     

Effects of water and nitrogen interaction on net photosynthesis, stomatal conductance, and water-use efficiency in two hybrid poplar clones

We examined the interactions of water and nitrogen availability by subjecting two Populus clones. Tristis and Eugenei, to five soil moisture and three soil nitrogen levels. Nitrogen application significantly increased net photosynthesis and stomatal conductance of flooded Eugenei and Tristis. The onset of flooding caused partial stomatal closure. Net photosynthesis significantly declined after a longer flooding period. Emergence of adventitious roots on the submerged portions of stems in both clones seemingly helped net photosynthesis fully recover in Eugenei and partially recover in Tristis. Under the progressive drought conditions, stomatal conductance was more sensitive to drought than net photosynthesis in both clones. Addition of nitrogen to progressively drying soil induced more stomatal closure in both clones. The highest water-use efficiency was found on the high-N/severe drought zone for Eugenei, whereas it was found on the high-N/mild to moderate drought zone for Tristis.     

Effects of flooding and drought on stomatal activity, transpiration, photosynthesis, water potential and water channel activity in strawberry stolons and leaves

Transpiration, xylem water potential and water channel activity were studied in developing stolons and leaves of strawberry (Fragaria × ananassa Duch.) subjected to drought or flooding, together with morphological studies of their stomata and other surface structures. Stolons had 0.12 stomata mm^–2 and a transpiration rate of 0.6 mmol H₂O m^–2 s^–1, while the leaves had 300 stomata mm^–2 and a transpiration rate of 5.6 mmol H₂O m^–2 s^–1. Midday water potentials of stolons were always less negative than in leaves enabling nutrient ion and water transport via or to the strawberry stolons. Drought stress, but not flooding, decreased stolon and leaf water potential from –0.7 to –1 MPa and from –1 to –2 MPa, respectively, with a concomitant reduction in stomatal conductance from 75 to 30 mmol H₂O m^–2 s^–1. However, leaf water potentials remained unchanged after flooding. Similarly, membrane vesicles derived from stolons of flooded strawberry plants showed no change in water channel activity. In these stolons, turgor may be preserved by maintaining root pressure, an electrochemical and ion gradient and xylem differentiation, assuming water channels remain open. By contrast, water channel activity was reduced in stolons of drought stressed strawberry plants. In every case, the effect of flooding on water relations of strawberry stolons and leaves was less pronounced than that of drought which cannot be explained by increased ABA. Stomatal closure under drought could be attributed to increased delivery of ABA from roots to the leaves. However, stomata closed more rapidly in leaves of flooded strawberry despite ABA delivery from the roots in the xylem to the leaves being strongly depressed. This stomatal closure under flooding may be due to release of stress ethylene. In the relative absence of stomata from the stolons, cellular (apoplastic) water transport in strawberry stolons was primarily driven by water channel activity with a gradient from the tip of the stolon to the base, concomitant with xylem differentiation and decreased water transport potential from the stolon tip to its base. Reduced water potential in the stolons under drought are discussed with respect to reduced putative water channel activity.     

Root signals and stomatal closure in relation to photosynthesis, chlorophyll a fluorescence and adventitious rooting of flooded tomato plants

Background and Aims

An investigation was carried out to determine whether stomatal closure in flooded tomato plants (Solanum lycopersicum) results from decreased leaf water potentials (ψ_L), decreased photosynthetic capacity and attendant increases in internal CO₂ (C_i) or from losses of root function such as cytokinin and gibberellin export.

Methods

Pot-grown plants were flooded when 1 month old. Leaf conductance was measured by diffusion porometry, the efficiency of photosystem II (PSII) was estimated by fluorimetry, and infrared gas analysis was used to determine C_i and related parameters.

Key Results

Flooding starting in the morning closed the stomata and increased ψ_L after a short-lived depression of ψ_L. The pattern of closure remained unchanged when ψ_`L depression was avoided by starting flooding at the end rather than at the start of the photoperiod. Raising external CO₂ concentrations by 100 µmol mol⁻¹ also closed stomata rapidly. Five chlorophyll fluorescence parameters [F_q′/F_m′, F_q′/F_v′, F_v′/F_m′, non-photochemical quenching (NPQ) and F_v/F_m] were affected by flooding within 12–36 h and changes were linked to decreased C_i. Closing stomata by applying abscisic acid or increasing external CO₂ substantially reproduced the effects of flooding on chlorophyll fluorescence. The presence of well-aerated adventitious roots partially inhibited stomatal closure of flooded plants. Allowing adventitious roots to form on plants flooded for >3 d promoted some stomatal re-opening. This effect of adventitious roots was not reproduced by foliar applications of benzyl adenine and gibberellic acid.

Conclusions

Stomata of flooded plants did not close in response to short-lived decreases in ψ_L or to increased C_i resulting from impaired PSII photochemistry. Instead, stomatal closure depressed C_i and this in turn largely explained subsequent changes in chlorophyll fluorescence parameters. Stomatal opening was promoted by the presence of well-aerated adventitious roots, implying that loss of function of root signalling contributes to closing of stomata during flooding. The possibility that this involves inhibition of cytokinin or gibberellin export was not well supported.Key words: Root to shoot communication, flooding stress, stomatal closure, photosynthesis, chlorophyll fluorescence, gas exchange, adventitious roots, plant hormones, abscisic acid, cytokinins, gibberellic acid     

Role of root systems of eastern larch and white spruce in response to flooding

Abstract. Soil flooding causes rapid reductions in transpiration, stomatal conductance and photosynthesis of many woody plants, which can decrease growth and ultimately result in plant death. This study was conducted to determine the role of the root system in the flooding response. Eastern larch ( Larix laricina ) seedlings were grown in Plexiglas tubes in which water uptake by flooded and unflooded roots was measured independently. Further flooding studies were conducted with eastern larch and white spruce ( Picea glauca ) in which stems were girdled. Root hydraulic properties were analysed using pressure-flow relationships. Transpiration rates of partially flooded plants declined more slowly than fully-flooded plants. Water uptake by unflooded roots of partially flooded seedlings increased momentarily with flooding. After lOd, flooding caused little change in root hydraulic conductance, a decrease in root system reflection coefficient, and an increase in osmotic permeability. Stem girdling had little effect on stomatal conductance and transpiration in comparison to flooding effects. The response of plant tops to flooding appears to be xylem-mediated and in proportion to the amount of root system flooded. Root hydraulic conductance appears to be unaffected by flooding except for a possible temporary increase on the first day following flooding treatments.     

Effect of spring flooding on endophyte differentiation,nitrogenase activity,root growth and shoot growth inMyrica gale

Summary Spring flooding was investigated as a possible limiting factor in the development of nitrogenase activity, root growth, and shoot growth inMyrica gale. Dormant, one year oldMyrica gale plants were placed in a greenhouse in early April and given three treatments: control (not flooded), flooded-water (flooded with water to 2.5 cm above the soil level) and flooded-peat (flooded with water-saturated peat to 4.0 cm above the soil level). Nitrogenase activity was absent at budbreak but appeared concurrently with the differentiation of vesicles by theFrankia sp. endophyte. Flooding delayed the onset of nitrogenase activity, substantially reduced the specific nitrogenase activity of the nodules, and also severely limited the production of the new nodule biomass. Consequently by 67 days past budbreak nitrogenase activity was much greater in the control plants (5.55±0.42 mol C₂H₄/plant.h; ± SE; N=9) than in the flooded-water (1.18±0.29) and flooded-peat (0.15±0.05) plants. Production of new secondary roots was substantially reduced in the flooded plants but adventitious roots were rapidly produced along the flooded portion of the stem in the better aerated zone near the surface. New nodules formed on several adventitious roots by 67 days indicating that the plants are able to replace their largely nonfunctional deeply flooded nodules with new nodules in the aerobic zone. Initially shoot growth was unaffected by flooding but by 67 days the flooded plants had substantially less leaf biomass, lower leaf and stem nitrogen concentrations, and less total shoot nitrogen content than the control plants.     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.     

Effect of Waterlogged Soil Conditions on the Production of Ethylene and on Water Relationships in Tomato Plants

The roots of tomato plants (Lycopersicon esculentum Mill., cv.Moneymaker) were exposed to low concentrations of oxygen bywaterlogging the soil or by growing the plants in nutrient solutionflushed with nitrogen gas. After 24 h, the rate of ethyleneproduction by the petioles, main stem, and shoot apex was increasedby 4–6-fold and the petioles developed epinastic curvatures.Removing the roots did not reproduce these responses. The amountsof ethylene produced by shoot tissues in response to physicalwounding was greatly increased by waterlogging the soil. The production of ethylene by roots was suppressed by the absenceof oxygen. When the roots were transferred back to an aerobicenvironment ethylene production quickly exceeded that observedin roots maintained continuously in aerobic conditions. The enhanced rate of ethylene production in the shoots occurredin the absence of increased water stress as measured with aleaf pressure chamber; leaf water potentials were increasedrather than decreased by waterlogging for 30 h or more. Thiswas associated with stomatal closure and reduced transpiration.Resistance to water flow through the plant increased as transpirationdecreased in response to waterlogging. However, at similar ratesof transpiration, resistance was normally lower in waterloggedplants than in controls.     

Antioxidative responses and morpho-anatomical adaptations to waterlogging in Sesbania virgata

Sesbania virgata (Leguminosae) is tolerant of long periods of soil inundation. However, its morphological adaptations to anoxia and its response to possible damage from oxidative stress are still unknown. Here, we provide new information that helps to explain the ability of S. virgata plants to grow in flooded environments. Plants containing six expanded leaves were placed in masonry tanks and were subjected to the following conditions: control (well watered), soil waterlogging (water to the setup level of 1 cm above the soil surface—roots and parts of the stems flooded), and complete submergence (whole plant flooded). Plants exposed to flooding (soil waterlogging and complete submergence) significantly increased their production of hydrogen peroxide (H₂O₂), indicating the extent of oxidative injury posed by stress conditions. We demonstrate that plants exposed to flooding develop an efficient scavenger of ROS (generated during stress) in the roots through the coordinated action of nonenzymatic ascorbic acid (Asc) and dehydroascorbate (DHA) as well as the enzymatic antioxidants superoxide dismutase (SOD), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), and glutathione reductase (GR) that are present in the tissues. Moreover, we observed the development of morpho-anatomical structures such as adventitious roots, lenticels, and cracks in the stem of plants under soil waterlogging. The secondary root of plants under soil waterlogging showed a thinner cortex and larger number of elements of small diameter vessels. Numerous aerenchymas were observed in the newly formed in the adventitious roots. We conclude that these antioxidative responses and morpho-anatomical adaptations in the roots are part of a suite of adaptations that allow S. virgata plants to survive long periods of flooding, notably under waterlogged conditions.     

Stem hypertrophic lenticels and secondary aerenchyma enable oxygen transport to roots of soybean in flooded soil

Background and Aims

Aerenchyma provides a low-resistance O₂ transport pathway that enhances plant survival during soil flooding. When in flooded soil, soybean produces aerenchyma and hypertrophic stem lenticels. The aims of this study were to investigate O₂ dynamics in stem aerenchyma and evaluate O₂ supply via stem lenticels to the roots of soybean during soil flooding.

Methods

Oxygen dynamics in aerenchymatous stems were investigated using Clark-type O₂ microelectrodes, and O₂ transport to roots was evaluated using stable-isotope ¹⁸O₂ as a tracer, for plants with shoots in air and roots in flooded sand or soil. Short-term experiments also assessed venting of CO₂ via the stem lenticels.

Key Results

The radial distribution of the O₂ partial pressure (pO₂) was stable at 17 kPa in the stem aerenchyma 15 mm below the water level, but rapidly declined to 8 kPa at 200–300 µm inside the stele. Complete submergence of the hypertrophic lenticels at the stem base, with the remainder of the shoot still in air, resulted in gradual declines in pO₂ in stem aerenchyma from 17·5 to 7·6 kPa at 13 mm below the water level, and from 14·7 to 6·1 kPa at 51 mm below the water level. Subsequently, re-exposure of the lenticels to air caused pO₂ to increase again to 14–17 kPa at both positions within 10 min. After introducing ¹⁸O₂ gas via the stem lenticels, significant ¹⁸O₂ enrichment in water extracted from roots after 3 h was confirmed, suggesting that transported O₂ sustained root respiration. In contrast, slight ¹⁸O₂ enrichment was detected 3 h after treatment of stems that lacked aerenchyma and lenticels. Moreover, aerenchyma accelerated venting of CO₂ from submerged tissues to the atmosphere.

Conclusions

Hypertrophic lenticels on the stem of soybean, just above the water surface, are entry points for O₂, and these connect to aerenchyma and enable O₂ transport into roots in flooded soil. Stems that develop aerenchyma thus serve as a ‘snorkel’ that enables O₂ movement from air to the submerged roots.     

Responses of red alder and black cottonwood seedlings to flooding

Red alder ( Alnus rubra Bong.) and black cottonwood ( Populus trichocarpa Torr. & Gray) seedlings were monitored to evaluate response during a 20-day period of artificial flooding and a 20-day recovery period following flooding. During the flooding period, both species showed changes in nutrient uptake and transport, initiated stemderived adventitious roots that became aerenchymatous, and exhibited hypertrophied lenticels. Flooded red alder seedlings also showed reduced height and leaf area growth and developed lower-stem hypertrophy. Flooded black cottonwood seedlings exhibited root dieback, aerenchyma in below ground root tips, and changes in root hydraulic conductance and xylem pressure potential. Contrary to expectations, however, stomatal closure following flooding was not observed in either species. Flooded red alder seedlings increased growth rapidly when drained, and by the end of the recovery period, formerly flooded and non-flooded red alder seedlings differed only minimally in this respect. In contrast, several characteristics of black cottonwood – including growth rate and nutrient content – still differed between formerly flooded and non-flooded seedlings at the end of the recovery period. Based on observed treatment differences at the end of the experiment, red alder seedlings were judged to be more tolerant of flooding than black cottonwood.     

Early stomatal closure in waterlogged pea plants is mediated by abscisic acid in the absence of foliar water deficits

Abstract Soil waterlogging decreased leaf conductance (interpreted as stomatal closure) of vegetative pea plants (Pisuin sativum L. cv. ‘Sprite’) approximately 24 h after the start of flooding, i.e. from the beginning of the second 16 h-long photo-period. Both adaxial and abaxial surfaces of leaves of various ages and the stipules were affected. Stomatal closure was sustained for at least 3 d with no decrease in foliar hydration measured as water content per unit area, leaf water potential or leaf water saturation deficit. Instead, leaves became increasingly hydrated in association with slower transpiration. These changes in the waterlogged plants over 3 d were accompanied by up to 10-fold increases in the concentration of endogenous abscisic acid (ABA). Waterlogging also increased foliar hydration and ABA concentrations in the dark. Leaves detached from non-waterlogged plants and maintained in vials of water for up to 3 d behaved in a similar way to leaves on flooded plants, i.e. stomata closed in the absence of a water deficit but in association with increased ABA content. Applying ABA through the transpiration stream to freshly detached leaflets partially closed stomata within 15 min. The extractable concentrations of ABA associated with this closure were similar to those found in flooded plants. When an ABA-deficient ‘wilty’ mutant of pea was waterlogged, the extent of stomatal closure was less pronounced than that in ordinary non-mutant plants, and the associated increase in foliar ABA was correspondingly smaller. Similarly, waterlogging closed stomata of tomato plants within 24 h, but no such closure was seen in ‘flacca’, a corresponding ABA-deficient mutant. The results provide an example of stomatal closure brought about by stress in the root environment in the absence of water deficiency. The correlative factor operating between the roots and shoots appeared to be an inhibition of ABA transport out of the shoots of flooded plants, causing the hormone to accumulate in the leaves.     

Stem hypoxia and root respiration of flooded maple and birch seedlings

Some researchers have attributed flood tolerance of woody plants to air entering the shoot through stems, leaves, or lenticels and diffusing to the roots to sustain aerobie respiration. The purpose of this study was to determine if internal aeration of roots by lower stems, changes in gross morphology of lower stems, or both, contribute to flood tolerance of certain tree species. Greenhouse-grown seedlings of red maple ( Acer rubrum L.) and river birch ( Betula nigra L.) tolerated at least 30 days of flooding, where as sugar maple ( Acer saccharum Marsh) and European white birch (also called silver birch, Betula pendula Roth) were intolerant. Flood treatment induced lentieel intumescences and adventitious root formation on red maple stems, but only adventitious roots formed on river birch stems. Stem morphology of sugar maple and European birch was unchanged by flooding. Flood stress decreased oxygen consumption capacity of excised roots from both tolerant and intolerant species. Exclusion of oxygen from the lower stems of flooded red maple and river birch prevented lenticel intumescence and adventitious root formation, but flood tolerance and root respiration capacity were unchanged. Neither internal aeration nor changes in stem morphology appear to account for flood tolerance of red maple and river birch.