Preference of wet dune species for waterlogged conditions can be explained by adaptations and specific recruitment requirements

The preference of wetland angiosperms for waterlogged soils has been explained by several hypotheses: (1) wetland species are adapted to waterlogging and sensitive to drought; (2) wetland species are tolerant to drought, but inferior competitors at drier conditions; (3) wetland species have narrow moisture optima for recruitment. We tested these hypotheses by the application of permanent and fluctuating water levels to experimental wet dune plant communities with four species that frequently occur in wet dune slacks (Carex flacca, Molinia caerulea, Samolus valerandi and Schoenus nigricans) and one competitive species of drier conditions (Calamagrostis epigejos). After 3 years, fluctuating water levels had led to lower total biomass production than permanent water levels, indicating that switching from aerated to anoxic soil conditions involved physiological costs. The collective biomass of wetland species was highest at permanently waterlogged conditions with interspecific variation in the biomass optima. At the nutrient-poor growth conditions of this long-term experiment, biomass of Calamagrostis was independent of water level treatment, thus the hypothesized competitive superiority of this species at drier conditions could not explain the biomass responses of the wetland species in this study. Instead, this is evidence for an ecological preference of adult wet dune plants for waterlogged to moist conditions. Recruitment of most wet dune slack species occurred in a narrower range of water levels than adult growth, indicating that recruitment requirements also pose a limitation to the distribution of these wet dune species. Incorporation of recruitment into nature management support models may improve their predictions.     

Growth and physiological responses of two mangrove species (Bruguiera gymnorrhiza and Kandelia candel) to waterlogging

Effects of duration of waterlogging on growth and physiological responses of two mangrove species, Bruguiera gymnorrhiza and Kandelia candel, were investigated. The relative growth rate of B. gymnorrhiza decreased significantly with waterlogged time, with the highest value found for drained plants and the lowest in plants under 12 weeks waterlogging. On the contrary, no significant difference was found between waterlogged and drained K. candel plants. The shoot to root biomass ratio of K. candel increased when subjected to 8 or 12 weeks waterlogging but little change was recorded in B. gymnorrhiza, indicating a shift in biomass allocation from roots to shoots in K. candel under prolonged waterlogging but not in B. gymnorrhiza. These different growth responses between the two mangrove species supported the hypothesis that K. candel is more tolerant to waterlogging than B. gymnorrhiza. Under 12 weeks waterlogged treatment, root oxidase activity significantly decreased in B. gymnorrhiza but increased in K. candel. Chlorophyll contents of K. candel increased more rapidly in response to waterlogging than B. gymnorrhiza. Activities of both peroxidase and superoxide dismutase increased significantly in leaves of K. candel when the waterlogging period was longer than 8 weeks, while only the peroxidase activity of B. gymnorrhiza showed a significant increase, indicating that K. candel had stronger resistance to the oxidant damage resulting from waterlogging. These physiological indicators further supported the hypothesis that K. candel is more tolerant to waterlogging than B. gymnorrhiza.     

Constraints of root response to waterlogging in Alisma triviale

To understand the economics of root aerenchyma formation in wetland plants, we investigated in detail the response of Alisma triviale to waterlogging. We hypothesized costs being associated with development of a large root air space. In three out-door pot experiments, seedlings (1 experiment) and mature plants (2 experiments) were grown under waterlogged and drained conditions for up to 2?months. Waterlogging promoted growth, and was associated with increased root porosity and decreased root density (fresh mass per volume). The increased formation of aerenchyma was associated with a higher root dry matter content for a given root density. Despite improved growth and earlier flowering, the waterlogged plants also showed signs of being constrained by the anoxic substrate, such as shallower roots, and a higher leaf dry matter content. The formation of aerenchyma was associated with costs, such as increased root dry matter content and reduced metaxylem vessel diameter. The faster growth of the seedlings under the waterlogged conditions, despite some signs of being stressed, was possibly a result of decreased requirements to allocate biomass below ground. In mature plants the increased aerenchyma allowed deeper root penetration, and ameliorated the effects of anoxia, reducing the differences in plant traits between the treatments.     

Inhibition of ethylene synthesis in tomato plants subjected to anaerobic root stress

Enhanced ethylene production and leaf epinasty are characteristic responses of tomato (Lycopersicon esculentum Mill.) to waterlogging. It has been proposed (Bradford, Yang 1980 Plant Physiol 65: 322-326) that this results from the synthesis of the immediate precursor of ethylene, 1-aminocyclopropane-1-carboxylic acid (ACC), in the waterlogged roots, its export in the transpiration stream to the shoot, and its rapid conversion to ethylene. Inhibitors of the ethylene biosynthetic pathway are available for further testing of this ACC transport hypothesis: aminooxyacetic acid (AOA) or aminoethoxyvinylglycine (AVG) block the synthesis of ACC, whereas CO²⁺ prevents its conversion to ethylene. AOA and AVG, supplied in the nutrient solution, were found to inhibit the synthesis and export of ACC from anaerobic roots, whereas Co²⁺ had no effect, as predicted from their respective sites of action. Transport of the inhibitors to the shoot was demonstrated by their ability to block wound ethylene synthesis in excised petioles. All three inhibitors reduced petiolar ethylene production and epinasty in anaerobically stressed tomato plants. With AOA and AVG, this was due to the prevention of ACC import from the roots as well as inhibition of ACC synthesis in the petioles. With Co²⁺, conversion of both root- and petiole-synthesized ACC to ethylene was blocked. Collectively, these data support the hypothesis that the export of ACC from low O₂ roots to the shoot is an important factor in the ethylene physiology of waterlogged tomato plants.     

Above-ground competition does not alter biomass allocated to roots in Abutilon theophrasti

We tested whether plants allocate proportionately less biomass to roots in response to above-ground competition as predicted by optimal partitioning theory. Two population densities of Abutilon theophrasti were achieved by planting one individual per pot and varying spacing among pots so that plants in the two densities experienced the same soil volume but different degrees of canopy overlap. Density did not affect root:shoot ratio, the partitioning of biomass between fine roots and storage roots, fine root length, or root specific length. Plants growing in high density exhibited typical above-ground responses to neighbours, having higher ratios of stem to leaf biomass and greater leaf specific area than those growing in low density. Total root biomass and shoot biomass were highly correlated. However, storage root biomass was more strongly correlated with shoot biomass than was fine-root biomass. Fine-root length was correlated with above-ground biomass only for the small subcanopy plants in crowded populations. Because leaf surface area increased with biomass, the ratio between absorptive root surface area and transpirational leaf surface area declined with plant size, a relationship that could make larger plants more susceptible to drought. We conclude that A. theophrasti does not reallocate biomass from roots to shoots in response to above-ground competition even though much root biomass is apparently involved in storage and not in resource acquisition.     

Waterlogging injury in sorghum seedlings and their kinetin effected rapid recovery

Short term waterlogging affected the growth of sorghum seedlings as indicated by a high mortality rate of the seedlings and a decrease in the shoot and root biomass, net assimilation rate (NAR), leaf area ratio (LAR), relative growth rate (RGR), total photosynthetic area and rate of photosynthesis. Plant resistance to the stress was exhibited by decreased stomatal index (SI), rate of transpiration and rate of respiration. When the stressed plants were exposed to normal conditions, the recovery of the seedlings was rapid and was enhanced by foliar application of kinetin at 5 mgl^-1.     

Growth,morphology and leaf characteristics after simulated herbivory in Chinese subtropical evergreen saplings

Growth, morphology and leaf characteristics were assessed in late spring following simulated autumnal defoliation in second-year saplings of three Chinese subtropical evergreen tree species.Castanopsis fargesii showed strong compensatory growth in terms of plant biomass after removal of both 50 and 75% of leaf biomass and slight compensatory growth after 90% defoliation. DefoliatedC. fargesii saplings had more leaves per unit shoot length than non-defoliated saplings. New leaves on defoliated plants were smaller and had higher per area nitrogen content than new leaves on non-defoliated plants.Pinus massoniana andElaeocarpus japonicus showed strong and no compensatory growth, respectively, after 50% defoliation. The strong compensatory growth inP. massoniana andC. fargesii may partly explain why these species predominate in the early and late successional phases of evergreen broad-leaved forests     

Escape from water or remain quiescent? Lotus tenuis changes its strategy depending on depth of submergence

Background and Aims

Two main strategies that allow plants to cope with soil waterlogging or deeper submergence are: (1) escaping by means of upward shoot elongation or (2) remaining quiescent underwater. This study investigates these strategies in Lotus tenuis, a forage legume of increasing importance in areas prone to soil waterlogging, shallow submergence or complete submergence.

Methods

Plants of L. tenuis were subjected for 30 d to well-drained (control), waterlogged (water-saturated soil), partially submerged (6 cm water depth) and completely submerged conditions. Plant responses assessed were tissue porosity, shoot number and length, biomass and utilization of water-soluble carbohydrates (WSCs) and starch in the crown.

Key Results

Lotus tenuis adjusted its strategy depending on the depth of submergence. Root growth of partially submerged plants ceased and carbon allocation prioritized shoot lengthening (32 cm vs. 24·5 cm under other treatments), without depleting carbohydrate reserves to sustain the faster growth. These plants also developed more shoot and root porosity. In contrast, completely submerged plants became quiescent, with no associated biomass accumulation, new shoot production or shoot elongation. In addition, tissue porosity was not enhanced. The survival of completely submerged plants is attributed to consumption of WSCs and starch reserves from crowns (concentrations 50–75 % less than in other treatments).

Conclusions

The forage legume L. tenuis has the flexibility either to escape from partial submergence by elongating its shoot more vigorously to avoid becoming totally submerged or to adopt a non-elongating quiescent strategy when completely immersed that is based on utilizing stored reserves. The possession of these alternative survival strategies helps to explain the success of L. tenuis in environments subjected to unpredictable flooding depths.     

The influence of some environmental factors on the production of Carex vesicaria and Phalaris arundinacea

The study deals with the annual cycles of biomass, productivity and necrosis of the perennial wetland species Carex vesicaria and Phalaris arundinacea, and with the main growth parameters during the first part of 1979. The amplitudes of biomass variation differ, but other features of the development are largely the same. To detect the various climatic factors which determine the above-ground biomass dynamics, a procedure of multiple stepwise regression has been used. Regressions for annual trend and variation in production were obtained. They provide coarse models, which may be improved by including translocation to belowground organs.Nomenclature follows P. Fournier (1977): Les quatres flores de France. Paris.     

An alternative mechanism for shade adaptation: implication of allometric responses of three submersed macrophytes to water depth

Allometric scaling models describing size-dependent biological relationships are important for understanding the adaptive responses of plants to environmental variation. In this study, allometric analysis was used to investigate the biomass allocation and morphology of three submerged macrophytes (Potamogeton maackianus, Potamogeton malaianus and Vallisneria natans) in response to water depth (1.0 and 2.5?m) in an in situ experiment. The three macrophytes exhibited different allometric strategies associated with distinct adjustments in morphology and biomass allocation in response to varying water depths. In deeper water, after accounting for the effects of plant size, P. maackianus and P. malaianus tended to enhance light harvesting by allocating more biomass to the stem, increasing shoot height and specific leaf area. V. natans tended to allocate more biomass to the leaf than to the basal stem (rosette), showing a higher leaf mass ratio and shoot height in deeper water. The three species decreased biomass allocation to roots as water depth increased. The main effect of water depth treatments was reduced light availability, which induced plastic shoot or leaf elongation. This shows that macrophytes have evolved responses to light limitation similar to those of terrestrial plants.     

Root morphology is related to the phenotypic variation in waterlogging tolerance of two populations of Suaeda salsa under salinity

The effects of waterlogging and salinity on seedling emergence, seedling growth and ion accumulation in a euhalophyte Suaeda salsa in an intertidal zone and on saline inland soil were investigated. Seedlings of S. salsa from the intertidal zone emerged more rapidly than those of the inland population under both waterlogged and drained conditions. Waterlogging and salinity had no adverse effects on seedling emergence of S. salsa from the intertidal zone, but markedly inhibited this parameter in the inland population. Waterlogging did not affect the seedling survival, shoot dry mass, and shoot height in high salinity in S. salsa from the intertidal zone, while the opposite trend was shown in the inland population. The root dry mass was higher in S. salsa from the intertidal zone as compared to the inland population, in waterlogged treatments by 1.9, 1.3, and 1.5 times in 1, 200, and 600 mM NaCl, respectively, and in drained treatments by 1.8, 2.3, and 3.0 times in 1, 200, and 600 mM NaCl, respectively. Waterlogging increased Na⁺ and K⁺ concentrations in high salinity, but waterlogging had no effect on Cl^- concentration in shoots of S. salsa from the intertidal zone. In all NaCl treatments, waterlogging had no effect on concentrations of these ions in shoots of S. salsa from the saline inland site. In a field investigation, the fresh mass of shoots and roots were lower, whereas the root/shoot ratio was 1.5 times higher in S. salsa from the intertidal zone, compared with the inland population. These findings indicate that S. salsa population from the intertidal zone is more waterlogging tolerant than the inland population. S. salsa from the intertidal zone produced relatively more root biomass and this might help anchor plants against tidal action in the intertidal zone. The physiological and morphological characteristics may determine the natural distributions of the two S. salsa populations in their different saline environments.     

Transpiration of a 64-year old maritime pine stand in Portugal

Alpine plant species have been shown to exhibit a more pronounced increase in leaf photosynthesis under elevated CO₂ than lowland plants. In order to test whether this higher carbon fixation efficiency will translate into increased biomass production under CO₂ enrichment we exposed plots of narrow alpine grassland (Swiss Central Alps, 2470 m) to ambient (355 μl l^-1) and elevated (680 μl l^-1) CO₂ concentration using open top chambers. Part of the plost received moderate mineral nutrient additions (40 kg ha^-1 year^-1 of nitrogen in a complete fertilizer mix). Under natural nutrient supply CO₂ enrichment had no effect on biomass production per unit land area during any of the three seasons studied so far. Correspondingly, the dominant species Carex curvula and Leontodon helveticus as well as Trifolium alpinum did not show a growth response either at the population level or at the shoot level. However, the subdominant generalistic species Poa alpina strongly increased shoot growth (+47%). Annual root production (in ingrowth cores) was significantly enhanced in C. curvula in the 2nd and 3rd year of investigation (+43%) but was not altered in the bulk samples for all species. Fertilizer addition generally stimulated above-ground (+48%) and below-ground (+26%) biomass production right from the beginning. Annual variations in weather conditions during summer also strongly influenced above-ground biomass production (19–27% more biomass in warm seasons compared to cool seasons). However, neither nutrient availability nor climate had a significant effect on the CO₂ response of the plants. Our results do not support the hypothesis that alpine plants, due to their higher carbon uptake efficiency, will increase biomass production under future atmospheric CO₂ enrichment, at least not in such late successional communities. However, as indicated by the response of P. alpina, species-specific responses occur which may lead to altered community structure and perhaps ecosystem functioning in the long-term. Our findings further suggest that possible climatic changes are likely to have a greater impact on plant growth in alpine environments than the direct stimulation of photosynthesis by CO₂. Counter-intuitively, our results suggest that even under moderate climate warming or enhanced atmospheric nitrogen deposition positive biomass responses to CO₂ enrichment of the currently dominating species are unlikely.     

Plant biomass and net production ofanogeissus latifolia Wall. in forests of semiarid zone of rajasthan (india)

A. latifolia grown in the Borimalan forest block in Prasad range (24°11′N and 73°42′ E) exerts clear positive correlations between CBH (circumference breast height)and number of growth rings of bole and branches, tree height, total biomass and leaf area. The net above-ground biomass is 3.95 × 10⁴ kg ha^-1. The average increment in non-photosynthetic (trunk + branch) biomass shows two peaks, the lower peak at 11–16 growth ring period, and the higher one at 34–36 growth ring period. The ratio of leaf dry weight/leaf area is16.3 to 34.8 mg cm^-2, the ratios between shoot net production: leaf weight and leaf area are1.5 g per g and 212 g m^-2 respectively.     

Effects of waterlogging on soil aeration and on root and shoot growth and yield of winter oats (Avena sativa L.)

Summary Winter oats were grown outdoors in lysimeters containing monoliths of a sandy loam soil. The soil was either freely-drained throughout the experiment or waterlogged to the soil surface from mid-January until mid-April. After the start of waterlogging the oxygen flux density decreased most rapidly nearer the soil surface and in the upper 50 cm declined to zero. At 80 cm depth the oxygen flux density at the end of the waterlogging still had not diminished to zero. While the soil was waterlogged root growth was negligible in the 20–50 cm zone of the soil profile, whereas below that depth root growth continued, reaching 95 cm by the end of the treatment. During the latter part of the waterlogging period root growth resumed in the upper 10 cm, and in the upper 2.5 cm was greater than in the freelydrained treatment.At the end of the waterlogging period, the total root length and shoot dry weights were 77 and 60% of those in the freely-drained treatment, tillering was restricted and leaf area index diminished. However, by anthesis, root length and shoot weights of the plants that had been waterlogged were only 10 and 12% less respectively than for the freely-drained plants. At harvest, total dry matter and grain yields were only 9% less, the latter largely through fewer grains per panicle.     

Mechanisms for the increase in phosphorus uptake of waterlogged plants: soil phosphorus availability, root morphology and uptake kinetics

Waterlogging frequently reduces plant biomass allocation to roots. This response may result in a variety of alterations in mineral nutrition, which range from a proportional lowering of whole-plant nutrient concentration as a result of unchanged uptake per unit of root biomass, to a maintenance of nutrient concentration by means of an increase in uptake per unit of root biomass. The first objective of this paper was to test these two alternative hypothetical responses. In a pot experiment, we evaluated how plant P concentration of Paspalum dilatatum, (a waterlogging-tolerant grass from the Flooding Pampa, Argentina) was affected by waterlogging and P supply and how this related to changes in root-shoot ratio. Under both soil P levels waterlogging reduced root-shoot ratios, but did not reduce P concentration. Thus, uptake of P per unit of root biomass increased under waterlogging. Our second objective was to test three non-exclusive hypotheses about potential mechanisms for this increase in P uptake. We hypothesized that the greater P uptake per unit of root biomass was a consequence of: (1) an increase in soil P availability induced by waterlogging; (2) a change in root morphology, and/or (3) an increase in the intrinsic uptake capacity of each unit of root biomass. To test these hypotheses we evaluated (1) changes in P availability induced by waterlogging; (2) specific root length of waterlogged and control plants, and (3) P uptake kinetics in excised roots from waterlogged and control plants. The results supported the three hypotheses. Soil P avail-ability was higher during waterlogging periods, roots of waterlogged plants showed a morphology more favorable to nutrient uptake (finer roots) and these roots showed a higher physiological capacity to absorb P. The results suggest that both soil and plant mechanisms contributed to compensate, in terms of P nutrition, for the reduction in allocation to root growth. The rapid transformation of the P uptake system is likely an advantage for plants inhabiting frequently flooded environments with low P fertility, like the Flooding Pampa. This advantage would be one of the reasons for the increased relative abundance of P. dilatatum in the community after waterlogging periods. Received: 15 February 1997 / Accepted: 20 May 1997     

The influence of water level on the growth and photosynthesis of Hydrocotyle ranunculoides L.fil.

Floating Pennywort (Hydrocotyle ranunculoides L.fil.), a native to North America and naturalized in Central and South America, is an invasive aquatic weed in western Europe and several other regions worldwide. H. ranunculoides settles primarily in stagnant to slow-flowing waters (e.g. ditches, canals, rivers, lakes and ponds). The species prefers sunny and nutrient-rich sites and forms dense interwoven mats, which can quickly cover the surface of infested waters. In this study, the effect of three different water levels on growth of Floating Pennywort was investigated. Plants were cultivated on high-nutrient soils under waterlogged, semi-drained and drained conditions. Highest relative growth rates (RGR) of 0.097±0.004 g g⁻¹ dw d⁻¹ were reached under waterlogged conditions. This was significantly higher than RGR of plants cultivated semi-drained and drained. Floating Pennywort showed some phenological adaptations to drained soil conditions, including significant differences in the relative amounts of leaf, petiole and shoot biomass, whilst the relative amount of root biomass was not significantly influenced by the water level. Furthermore, Floating Pennywort reached under drained conditions lower relative water contents (RWC) of leaves, petioles and shoots, a significant shorter length of internodes, a significant lower extent of shoot porosity (POR), a lower chlorophyll content and an increased Chl_a:Chl_b ratio. In addition, maximum gas exchange of drained cultivated plants is significantly lower, due to strongly decreased leaf conductance under reduced water availability. Overall, H. ranunculoides showed ability to grow under different water levels, but performed best under waterlogged conditions.     

A Model Allocating Growth Among Leaf Proteins, Shoot Structure, and Root Biomass to Produce Balanced Activity

A model is developed that considers the allocation of carbonand nitrogen substrates to a protein compartment in the shoots,shoot structural components, and root biomass. Inclusion ofa shoot-protein compartment allows variation in shoot-specificactivity to be modelled as a function of leaf nitrogen concentration.Allocation to the biomass compartments is controlled by twopartitioning variables that are defined by explicitly usingthe balanced activity hypothesis. The model produces balancedactivity where the shoot-specific activity, as well as rootand shoot biomass, vary in response to the above-ground (lightand CO₂) and below-ground (nitrogen) environments. The predictedpatterns of both root: shoot ratio and leaf nitrogen concentrationin response to environmental resource availability are qualitativelyconsistent with general trends observed in plants. Biomass allocation, plant growth, modelling, leaf nitrogen, root: shoot ratio, balanced activity     

Review of wheat improvement for waterlogging tolerance in Australia and India: the importance of anaerobiosis and element toxicities associated with different soils

Background and Aims

The lack of knowledge about key traits in field environments is a major constraint to germplasm improvement and crop management because waterlogging-prone environments are highly diverse and complex, and the mechanisms of tolerance to waterlogging include a large range of traits. A model is proposed that waterlogging tolerance is a product of tolerance to anaerobiosis and high microelement concentrations. This is further evaluated with the aim of prioritizing traits required for waterlogging tolerance of wheat in the field.

Methods

Waterlogging tolerance mechanisms of wheat are evaluated in a range of diverse environments through a review of past research in Australia and India; this includes selected soils and plant data, including plant growth under waterlogged and drained conditions in different environments. Measurements focus on changes in redox potential and concentrations of diverse elements in soils and plants during waterlogging.

Key Results

(a) Waterlogging tolerance of wheat in one location often does not relate to another, and (b) element toxicities are often a major constraint in waterlogged environments. Important element toxicities in different soils during waterlogging include Mn, Fe, Na, Al and B. This is the first time that Al and B toxicities have been indicated for wheat in waterlogged soils in India. These results support and extend the well-known interactions of salinity/Na and waterlogging/hypoxia tolerance.

Conclusions

Diverse element toxicities (or deficiencies) that are exacerbated during waterlogging are proposed as a major reason why waterlogging tolerance at one site is often not replicated at another. Recommendations for germplasm improvement for waterlogging tolerance include use of inductively coupled plasma analyses of soils and plants.Key words: Waterlogging, microelements, toxicity, redox potential, wheat, anaerobiosis

‘No grain is ever produced without water, but too much water tends to spoil the grain and inundation is as injurious to growth as dearth of water.’ Narada Smriti XI, 19; circa 3000 bc.‘Waterlogging’ is defined as a condition of the soil where excess water limits gas diffusion; while ‘waterlogging tolerance’ is defined as survival or the maintenance of high growth rates, biomass accumulation or grain yield under waterlogging relative to non waterlogged (usually drained soil) conditions (Setter and Waters, 2003).

     

Plant growth, nutrient acquisition and mycorrhizal symbioses of a waterlogging tolerant legume (Lotus glaber Mill.) in a saline-sodic soil

Seedlings of Lotus glaberMill., were grown in a native saline-sodic soil in a greenhouse for 50 days and then subjected to waterlogging for an additional period of 40 days. The effect of soil waterlogging was evaluated by measuring plant growth allocation, mineral nutrition and soil chemical properties. Rhizobiumnodules and mycorrhizal colonisation in L. glaberroots were measured before and after waterlogging. Compared to control plants, waterlogged plants had decreased root/shoot ratio, lower number of stems per plant, lower specific root length and less allocation of P and N to roots. Waterlogged plants showed increased N and P concentrations in plant tissues, larger root crown diameter and longer internodes. Available N and P and organic P, pH and amorphous iron increased in waterlogged soil, but total N, EC and exchangeable sodium were not changed. Soil waterlogging decreased root length colonised by arbuscular mycorrhizal (AM) fungi, arbuscular colonisation and number of entry points per unit of root length colonised. Waterlogging also increased vesicle colonisation and Rhizobium nodules on roots. AM fungal spore density was lower at the end of the experiment in non-waterlogged soil but was not reduced under waterlogging. The results indicate that L. glaber can grow, become nodulated by Rhizobium and colonised by mycorrhizas under waterlogged condition. The responses of L. glaber may be related its ability to form aerenchyma.