An overview of oxygen transport in plants: diffusion and convection.

The movement of gases within plants is crucial for species that live in flood-prone areas with limited soil oxygen. These plants adapt to hypoxia/anoxia not by using oxygen more efficiently, but by ensuring a steady oxygen supply to their cells. Wetland plants typically form gas-filled spaces (aerenchyma) in their tissues, providing a low-resistance pathway for gas movement between shoots and roots, especially when the shoots are above water, and the roots are submerged. Oxygen movement in plant roots is mainly through diffusion. However, in certain species, such as emergent and floating-leaved plants, pressurized flows can also facilitate the movement of gases within their stems and rhizomes. Three types of pressurized (convective) flows have been identified: humidity-induced pressurization (positive pressure), thermal osmosis (positive pressure with air flow against the heat gradient), and venturi-induced suction (negative pressure) caused by wind passing over broken culms. A clear diel variation in pressurized flows exists, with higher pressures and flows during the day and negligible pressures and flows during the night. This article discusses some key aspects of these mechanisms for oxygen movement.

Echinochloa crus-galli seed physiological dormancy and germination responses to hypoxic floodwaters.

Hypoxic floodwaters can seriously damage seedlings. Seed dormancy could be an effective trait to avoid lethal underwater germination. This research aimed to discover novel adaptive dormancy responses to hypoxic floodwaters in seeds of Echinochloa crus-galli, a noxious weed from rice fields and lowland croplands. Echinochloa crus-galli dormant seeds were subjected to a series of sequential treatments. Seeds were: (i) submerged under hypoxic floodwater (simulated with hypoxic flasks) at different temperatures for 15 or 30 days, and germination tested under drained conditions while exposing seeds to dormancy-breaking signals (alternating temperatures, nitrate (KNO3 ), light); or (ii) exposed to dormancy-breaking signals during hypoxic submergence, and germination monitored during incubation and after transfer to drained conditions. Echinochloa crus-galli seed primary dormancy was attenuated under hypoxic submergence but to a lesser extent than under drained conditions. Hypoxic floodwater did not reinforced dormancy but hindered secondary dormancy induction in warm temperatures. Seeds did not germinate under hypoxic submergence even when subjected to dormancy-breaking signals; however, these signals broke dormancy in seeds submerged under normoxic water. Seeds submerged in hypoxic water could sense light through phytochrome signals and germinated when normoxic conditions were regained. Hypoxic floodwaters interfere with E. crus-galli seed seasonal dormancy changes. Dormancy-breaking signals are overridden during hypoxic floods, drastically decreasing underwater germination. In addition, results indicate that a fraction of E. crus-galli seeds perceive dormancy-breaking signals under hypoxic water and germinate immediately after aerobic conditions are regained, a hazardous yet less competitive environment for establishment.

Different strategies of Lotus japonicus, L. corniculatus and L. tenuis to deal with complete submergence at seedling stage.

Two main strategies allow plants to deal with submergence: (i) escape from below water by means of shoot elongation, or (ii) remaining quiescent under the water until water subsides and then resume growth. We investigated these strategies in seedlings of Lotus japonicus, L. corniculatus and L. tenuis subjected to control and submergence for 12 days, with a subsequent 30-day recovery period. All three species survived submergence but used different strategies. Submerged seedlings of L. japonicus exhibited an escape strategy (emerging from water) as a result of preferential carbon allocation towards shoot mass and lengthening, in detriment to root growth. In contrast, seedlings of L. corniculatus and L. tenuis became quiescent, with no biomass accumulation, no new unfolding of leaves and no shoot elongation. Upon de-submergence, seedlings of L. japonicus had the lowest recovery growth (a biomass and shoot height 58% and 40% less than controls, respectively), L. corniculatus was intermediate and L. tenuis showed the greatest recovery growth. Previously submerged seedlings of L. tenuis did not differ from their controls, either in final shoot biomass or shoot height. Thus, for the studied species, quiescence appears to be an adequate strategy for tolerance of short-term (i.e., 12 days) complete submergence, being consistent with field observations of L. tenuis colonisation of flood-prone environments.

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.

Flooding effects on plants recovering from defoliation in Paspalum dilatatum and Lotus tenuis.

BACKGROUND AND AIMS: Flooding and grazing are major disturbances that simultaneously affect plant performance in many humid grassland ecosystems. The effects of flooding on plant recovery from defoliation were studied in two species: the grass Paspalum dilatatum, regrowing primarily from current assimilation; and the legume, Lotus tenuis, which can use crown reserves during regrowth. METHODS: Plants of both species were subjected to intense defoliation in combination with 15 d of flooding at 6 cm water depth. Plant recovery was evaluated during a subsequent 30-d growth period under well-watered conditions. Plant responses in tissue porosity, height, tiller or shoot number and biomass of the different organs were assessed. KEY RESULTS: Flooding increased porosity in both P. dilatatum and L. tenuis, as expected in flood-tolerant species. In P. dilatatum, defoliation of flooded plants induced a reduction in plant height, thus encouraging the prostrated-growth response typical of defoliated plants rather than the restoration of contact with atmospheric oxygen, and most tillers remained submerged until the end of the flooding period. In contrast, in L. tenuis, plant height was not reduced when defoliated and flooded, a high proportion of shoots being presented emerging above water (72 %). In consequence, flooding plus defoliation did not depress plant recovery from defoliation in the legume species, which showed high sprouting and use of crown biomass during regrowth, whereas in the grass species it negatively affected plant recovery, achieving 32 % lower biomass than plants subjected to flooding or defoliation as single treatments. CONCLUSIONS: The interactive effect of flooding and defoliation determines a reduction in the regrowth of P. dilatatum that was not detected in L. tenuis. In the legume, the use of crown reserves seems to be a key factor in plant recovery from defoliation under flooding conditions.

Trade-off between root porosity and mechanical strength in species with different types of aerenchyma.

The objective of this work was to study the existence of a trade-off between aerenchyma formation and root mechanical strength. To this end, relationships among root anatomical traits and mechanical properties were analysed in plant species with contrasting root structural types: Paspalidium geminatum (graminaceous type), Cyperus eragrostis (cyperaceous type), Rumex crispus (Rumex type) and Plantago lanceolata (Apium type). Variations in anatomical traits and mechanical strength were assessed as a function of root diameter by exposing plants to 0, 7, 15 and 30 d of control and flooded conditions. For each species, the proportion of root cortex was positively associated with the increment of root diameter, contributing to the increase in root porosity under both control and flooded conditions. Moreover, cell lysis produced an additional increase in root porosity in most species under flooded conditions (except R. crispus). Both structural types that presented a uniseriate layer (epidermis) to cope with compression (Rumex and Apium types) were progressively weakened as root porosity increased. This effect was significant even when the increment of root porosity was solely because of increased root diameter (R. crispus), as when both processes (root diameter and cell lysis) added porosity to the roots (P. lanceolata). Conversely, structural types that presented a multiseriate ring of cells in the outer cortex (graminaceous and cyperaceous types) maintained mechanical strength over the whole range of porosity, in spite of lysogenic processes registered in the inner cortex. In conclusion, our study demonstrates a strong trade-off between aerenchyma formation and mechanical strength in root structural types that lacked a multiseriate ring of tissue for mechanical protection in the outer cortex. The results suggest that this ring of tissue plays a significant role in maintaining the mechanical strength of roots when flooding induces the generation of additional aerenchyma tissue in the root cortex.