Oxygen absorption by adventitious roots promotes the survival of completely submerged terrestrial plants.

Background and Aims Flooding imposes stress upon terrestrial plants because it results in oxygen deficiency, which is considered a major problem for submerged plants. A common response of terrestrial plants to flooding is the formation of aquatic adventitious roots. Some studies have shown that adventitious roots on submerged plants are capable of absorbing water and nutrients. However, there is no experimental evidence for the possible oxygen uptake function of adventitious roots or for how important this function might be for the survival of plants during prolonged submergence. This study aims to investigate whether adventitious roots absorb oxygen from the water column, and whether this new function is beneficial to the survival of completely submerged plants. Methods Taking Alternanthera philoxeroides (Mart.) Griseb. as a representative species, the profiling of the underwater oxygen gradient towards living and dead adventitious roots on completely submerged plants was conducted, the oxygen concentration in stem nodes with and without adventitious roots was measured, and the growth, survival and non-structural carbohydrate content of completely submerged plants with and without adventitious roots was investigated. Key Results Oxygen profiles in the water column of adventitious roots showed that adventitious roots absorbed oxygen from water. It is found that the oxygen concentration in stem nodes having adventitious roots was higher than that in stem nodes without adventitious roots, which implies that the oxygen absorbed by adventitious roots from water was subsequently transported from the roots to other plant tissues. Compared with plants whose adventitious roots had been pruned, those with intact adventitious roots had slower leaf shedding, slower plant mass reduction, more efficient carbohydrate economy and prolonged survival when completely submerged. Conclusions The adventitious roots of A. philoxeroides formed upon submergence can absorb oxygen from ambient water, thereby alleviating the adverse effects of oxygen deficiency, enabling efficient utilization of carbohydrates and delaying the death of completely submerged plants.

Water depth-dependent stem elongation of completely submerged Alternanthera philoxeroides is mediated by intra-internodal growth variations.

Complete submergence, especially deep submergence, poses a serious threat to the growth and survival of plants. One study previously showed that Alternanthera philoxeroides (a herbaceous perennial plant) submerged at depth of 2 m presented fast stem elongation and reduced stem elongation as water depth increased. In the present study, we aimed to figure out from the morphological and anatomical perspective how the differential growth response of the plant to water depth was achieved. We investigated the elongation of different stem parts and the relationship of stem elongation to cell size and number in A. philoxeroides by conducting experiments using a series of submergence depths (0 m, 2 m, 5 m, and 9 m). The results showed that, in comparison with unsubmerged plants, completely submerged plants exhibited enhanced elongation at depths of 2 m and 5 m but suppressed elongation at depth of 9 m in immature stem internodes, and displayed very little elongation in mature stem internodes at any depths. The stem growth of A. philoxeroides at any submergence depth was chiefly caused by the elongation of the basal parts of immature internodes. The elongation of the basal parts of immature internodes was highly correlated to both cell proliferation and cell enlargement, but the elongation of the middle and upper parts of immature internodes correlated nearly only with cell enlargement. This study provided new information on the growth responses of A. philoxeroides to heterogeneous submergence environments and deepened our understanding of the growth performance of terrestrial plants in habitats prone to deep floods.

Stem elongation and gibberellin response to submergence depth in clonal plant Alternanthera philoxeroides.

Clonal plants are widely distributed in the riparian zone and play a very important role in the maintenance of wetland ecosystem function. Flooding is an environmental stress for plants in the riparian zone, and the response of plants varies according to the depth and duration of flooding. However, there is a lack of research on the growth response of clonal plants during flooding, and the endogenous hormone response mechanism of clonal plants is still unclear. In the present study, Alternanthera philoxeroides, a clonal plant in the riparian zone, was used to investigate the time-dependent stem elongation, the elongation of different part of the immature internodes, and the relationship between growth elongation and the phytohormone gibberellin (GA) under a series of submergence depths (0 m, 2 m, 5 m, and 9 m). The results showed that stem elongation occurred under all treatments, however, compared to 0 m (control), plants grew more under 2 m and 5 m submergence depth, while grew less under 9 m water depth. Additionally, basal part elongation of the immature internode was the predominant factor contributing to the stem growth of A. philoxeroides under different submergence depths. The phytohormone contents in basal part of the mature and immature internodes showed that GA induced the differential elongation of internode. Plant submerged at depth of 2 m had the highest GA accumulation, but plant submerged at depth of 9 m had the lowest GA concentration. These data suggested that GA biosynthesis are essential for stem elongation in A. philoxeroides, and the basal part of the immature internode was the main position of the GA biosynthesis. This study provided new information about the rapid growth and invasion of the clonal plant A. philoxeroides around the world, further clarified the effects of submergence depth and duration on the elongation of the stem, and deepened our understanding of the growth response of terrestrial plants in deeply flooded environments.

Differential Growth Responses of Alternanthera philoxeroides as Affected by Submergence Depths.

Global climate change has resulted in an increase in intensity and frequency of flooding, plants living in lowlands, and shore areas have to confront submergence caused by flooding, submergence-tolerant plants usually respond by adopting either escape or quiescence strategies. While certain plants exhibit a changeover from escape strategy upon partial submergence to quiescence strategy under complete shallow submergence, it remains unknown whether plants completely submerged at different water depths would adjust their strategies to cope with the change in submergence depth. Alternanthera philoxeroides is an ideal species to explore this adjustment as it is widely distributed in flood-disturbed habitats and exhibits an escape strategy when completely submerged in shallow waters. We investigated the responses of A. philoxeroides in terms of morphology, anatomy, and non-structural carbohydrate metabolism by conducting experiments using a series of submergence depths (0, 2, 5, and 9 m). During the submergence treatment, environmental factors such as light, dissolved oxygen, and temperature for submerged plants were kept constant. The results showed that A. philoxeroides plants submerged at depth of 2 m presented an escape strategy via fast stem elongation, extensive pith cavity development, and small biomass loss. However, the retarded stem elongation, reduced pith cavity transverse area, and increased biomass loss along the water depth gradient indicated that A. philoxeroides altered its growth response as water depth increased from 2 to 9 m. It is found that the changeover of response strategies occurred at higher submergence depths (5-9 m). Based on the results of our experiments, we demonstrated that water depth played an important role in driving the change in strategy. The water-depth-dependent growth performance of A. philoxeroides would benefit the species in habit exploration and exploitation. Further studies should focus on the performances of plants when submerged at varied water depths with different light climates and dissolved oxygen content, and how water depths drive the response behaviors of the submerged plants.

Similar Growth Performance but Contrasting Biomass Allocation of Root-Flooded Terrestrial Plant Alternanthera philoxeroides (Mart.) Griseb. in Response to Nutrient Versus Dissolved Oxygen Stress.

Terrestrial plants may experience nutrient and oxygen stress when they are submerged, and increases in flooding are anticipated with climate change. It has been well reported that plants usually shift biomass allocation and produce more roots in response to nutrient deficiency. However, it is unclear whether plants experiencing oxygen deficiency stimulate biomass allocation to roots to enhance nutrient absorption, similar to how plants experiencing nutrient deficiency behave. We investigated the responses of the terrestrial species Alternanthera philoxeroides, upon root flooding, to nutrient versus dissolved oxygen deficiency in terms of plant growth, biomass allocation, root production, root efficiency (plant growth sustained per unit root surface area), and root aerenchyma formation. Both nutrient and dissolved oxygen deficiency hampered the growth of root-flooded plants. As expected, plants experiencing nutrient deficiency increased biomass allocation to roots and exhibited lower root efficiency; in contrast, plants experiencing dissolved oxygen deficiency decreased biomass allocation to roots but achieved higher root efficiency. The diameter of aerenchyma channels in roots were enlarged in plants experiencing dissolved oxygen deficiency but did not change in plants experiencing nutrient deficiency. The widening of aerenchyma channels in roots could have improved the oxygen status and thereby the nutrient absorption capability of roots in low oxygen environments, which might benefit the plants to tolerate flooding.

Publisher Correction: Effects of water level regulation on the seed germination and production of annual plant Xanthium sibiricum in the water-level-fluctuating-zone of Three Gorges Reservoir.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Effects of water level regulation on the seed germination and production of annual plant Xanthium sibiricum in the water-level-fluctuating-zone of Three Gorges Reservoir.

Vegetation effectively slows down the deterioration rate of the ecosystem in the water-level fluctuation zone (WLFZ). In this study, we investigated the impact of impoundment on the germination of seeds from Xanthium sibiricum fruits with various degrees of maturity and produced by X. sibiricum plants growing at different elevations in the WLFZ. Seed production by X. sibiricum under the current water level regulation mode was also investigated. Seeds were collected from seven elevations of the WLFZ of Three Gorges Reservoir (TGR) and exposed at these elevations to submergence. Seed production of the plants germinated from X. sibiricum seeds was observed. The maturity degree of X. sibiricum fruits from different elevations has no influence on the seed production by the plants that are produced from the fruits. Under the present water-level regulation mode, X. sibiricum growing above 155 m are able to generate mature seeds and thus provide seed sources for population maintenance, although the plants growing at an elevation below 165 m failed to produce fruits in 2010 due to unusual flooding. This study is useful for the vegetation recovery and reconstruction in other regions with hydrological characteristics similar to that of the TGR.

Seed sojourn and fast viability loss constrain seedling production of a prominent riparian protection plant Salix variegata Franch.

Salix variegata Franch, a prominent plant applied in riparian shelter vegetation in Three Gorges reservoir region of China, produces many seeds every year but generates only a few or no seedlings. Whether the low seedling production of S. variegata is caused by seed sterility or by rapid loss of seed viability remains unknown. We investigated the sojourn time of mature seeds in capsules produced in early, mid, and late reproductive season and the germinability of mature seeds fresh or stored after different period of time. The sojourn time of seeds in capsules was 2.89, 3.95, and 4.72 days in early, mid, and late reproductive season, respectively. The maximal germination percentage of non-stored fresh seeds produced in early, mid, and late reproductive season was 93.33%, 78.67%, and 40%, respectively, which indicates mature seeds were not sterile. The longest viability-retaining time of seeds produced in early, mid, and late reproductive season was only 8, 16, 16 days, respectively, indicating that mature seeds of S. variegata lost viability very rapidly. Mature seeds possessed good viability, but their rapid viability loss caused the low seedling production and hampered the population growth of S. variegata in the riparian area of Three Gorges reservoir region.