Water relations,ethylene production,and morphological adaptation ofFraxinus pennsylvanica seedlings to flooding

Summary Fraxinus pennsylvanica Marsh. seedlings that were 150 days old adapted well to flooding of soil with stagnant water for 28 days. Early stomatal closure, followed by reopening as well as hypertrophy of lenticels and formation of adventitious roots on submerged portions of stems appeared to be important adaptations for flood tolerance. Leaf water potential (₁) was consistently higher in flooded than in unflooded seedlings, indicating higher leaf turgidity in the former. This was the result of (1) early reduction in transpiration associated with stomatal closure, and (2) subsequently increased absorption of water by the newly-formed adventitious roots as stomata reopened and transpiration increased. Waterlogging of soil was followed by large increases in ethylene content of stems, both below and above the level of submersion. Formation of hypertrophied lenticels and adventitious roots on flooded plants was correlated with increased ethylene production. However, the involvement of various compounds other than ethylene in inducing morphological changes in flooded plants is also emphasized.Research supported by the College of Agricultural and Life Sciences, University of Wisconsin, Madison, WI, USA     

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.     

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.     

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

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

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.     

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.     

Responses of Melaleuca quinquenervia seedlings to flooding

Abstract Studies were conducted on effects of flooding for 15, 30, 60, and 90 days on morphological changes, stomatal aperture, water potential, and growth of seedlings of Melaleuca quinquenervia, a species often planted for reclamation of swamps. Flooding rapidly induced formation of many hair-like adventitious roots as well as a few thick adventitious roots that originated on the original root system. Some adventitious roots also formed on submerged portions of the stem. Melaleuca seedlings were very tolerant of flooding as shown by only slight reduction in dry weight increment of shoots after 30 days of flooding in stagnant water. Although flooding for 60 or 90 days significantly reduced dry weight increment of leaves, dry weight increment of roots was not inhibited by any flooding treatment, reflecting both degeneration of some of the original roots and compensatory growth of adventitious roots. On certain days flooding induced stomatal closure on both adaxial and abaxial leaf surfaces. Extensive production of adventitious roots and some stomatal reopening after a critical period of flooding appeared to be important factors in the flooding tolerance of Melaleuca and are consistent with its aggressiveness and vigorous growth on wet sites.     

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.     

淹水对玉米不定根形态结构和ATP酶活性的影响

淹水２天后,玉米苗基节内即有不定根原基一进于正常植株。淹水１６天后,从基节部长出的不定根数多于正常植株,但淹水导致根系生长和干物质积累大幅度下降。淹水幼苗不定根伸长区内有发达的通气组织形成,使根内部组织孔隙度大幅提高。电镜细胞化学研究表明,经１５天淹一,不定根根尖细胞内ＡＴＰ酶的分布与正常功苗基本相同,酶活性尽管有一定的下降,但仍保持较高水平。根据实验结果,本文重点讨论了不定根的发生及其内部通气组     

水翁(Cleistocalyx operculatus)幼苗对淹水的反应初报

研究了水翁在生理和形态上对3个月淹水期的反应。在潮湿或淹水条件下水翁能存活并保持一定的净光合速率和生长速率,全淹条件下存活期为60d,水翁对淹水的适应包括：（1）淹和的茎部产生肥大皮孔和不定根,（2）不定根系的活力比正常根系的活力高,有不定根的植株的气孔传导率和蒸腾速率比没有不定根的植株高得多。水翁是一种耐淹植物,可在河岸、库岸等水位波动地区种植。     

Ecophysiological adaptations of black spruce (<Emphasis Type="Italic"> Picea mariana</Emphasis>) and tamarack (<Emphasis Type="Italic"> Larix laricina</Emphasis>) seedlings to flooding

Black spruce [ Picea mariana (Mill.) B.S.P.] and tamarack [ Larix laricina (Du Roi) K. Koch] are the predominant tree species in boreal peatlands. The effects of 34 days of flooding on morphological and physiological responses were investigated in the greenhouse for black spruce and tamarack seedlings in their second growing season (18 months old). Flooding resulted in reduced root hydraulic conductance, net assimilation rate and stomatal conductance and increased needle electrolyte leakage in both species. Flooded tamarack seedlings maintained a higher net assimilation rate and stomatal conductance compared to flooded black spruce. Flooded tamarack seedlings were also able to maintain higher root hydraulic conductance compared to flooded black spruce seedlings at a comparable time period of flooding. Root respiration declined in both species under flooding. Sugar concentration increased in shoots while decreasing in roots in both species under flooding. Needles of flooded black spruce appeared necrotic and electrolyte leakage increased over time with flooding and remained significantly higher than in flooded tamarack seedlings. No visible damage symptoms were observed in flooded tamarack seedlings. Flooded tamarack seedlings developed adventitious roots beginning 16 days after the start of flooding treatment. Adventitious roots exhibited significantly higher root hydraulic conductivity than similarly sized flooded tamarack roots. Flooded black spruce lacked any such morphological adaptation. These results suggest that tamarack is better able to adjust both morphologically and physiologically to prolonged soil flooding than black spruce seedlings.     

EFFECTS OF FLOODING AND NUTRIENT ENRICHMENT ON BIOMASS ALLOCATION IN ACER RUBRUM SEEDLINGS

A greenhouse experiment was conducted on Acer rubrum seedlings to evaluate the effects of flood frequency on production and allocation of biomass and to test the effects of N and P fertilization on production and allocation. Seedlings from the Dismal Swamp were subjected to three flood treatments (no flooding, intermittent flooding, and continuous flooding) and four enrichment treatments (no enrichment, N additions, P additions, and N + P additions). More continuous flooding resulted in less biomass production. Biomass increased during the study in all treatments except for root mass in the continuously flooded treatment. However, production of abundant adventitious roots compensated for the lack of normal root growth. Root/shoot ratios exhibited the greatest decreases in the continuously flooded plants. Plants with N + P added had significantly more leaf, stem, and total mass than the nonenriched plants four months into the study. The N + P additions had apparently compensated for the effects of flood stress in the continuously flooded plants by the end of the study. The fertilized seedlings accumulated higher concentrations of N and P, but their nutrient use efficiency (biomass production per unit nutrient absorbed) was lower than in the nonenriched plants. Acer rubrum seedlings survive flooded conditions through several adaptations; however, theirgrowth is slowed by continuous flooding.     

Effect of flooding and temperature regime on growth and stomatal resistance ofBetula platyphylla var.japonica seedlings

Summary Both flooding and low temperature reduced height and stem diameter growth; leaf initiation; growth of leaves, stems, and roots; and lowered root-shoot ratios of 112-dayoldBetula platyphylla var.japonica seedlings. Flooding also induced leaf scorching and abscission. Growth was reduced much more by flooding than by low temperature. Interactive effects of flooding and temperature were shown on height growth, leaf initiation and expansion, and dry weight increment of leaves, stems and roots. The amount of growth reduction by flooding and low temperature was greater when based on analysis of dry weight increment of leaves, stems, and roots, than on their relative growth rates. The greater reduction of growth by flooding than by low temperature was associated with fewer and smaller leaves, more leaf injury, more stomatal closure, and greater decay of roots in flooded plants. Flooding and low temperature appeared to reduce growth by somewhat different physiological mechanisms. Research supported by the College of Agricultural and Life Sciences, University of Wisconsin, Madison, WI, USA and by Yamagata University, Tsuruoka, Japan. McIntyre-Stennis Project 2599.     

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

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

Physiological and morphological responses of red alder and sitka alder to flooding

Red alder (Alnus rubra Bong.) and sitka alder (A. viridis ssp. sinuata [Regel] Löve & Löve) are nitrogen-fixing woody species that grow sympatrically along the Pacific coast of North America. Red alder is found in poorly drained lowlands, as well as in soils of moist upland slopes, whereas sitka alder generally colonizes well-drained soils. To identify factors that contribute to flood tolerance, we conducted greenhouse experiments subjecting both species to a 20-day flood and 10-day recovery and red alder to a 50-day flood and 20-day recovery. We determined the effect of this stress on nitrogenase activity, root and nodule alcohol dehydrogenase (ADH) activity, lenticel and adventitious root development, relative growth rate (RGR), and leaf gas exchange. After 24 h of flooding, nitrogenase activity could not be detected in either species. Limited nitrogenase activity did return in red alder at the end of a 10-day recovery following the 20-day flood, but sitka alder showed no recovery of nitrogenase activity. After 50 days of continuous flooding, red alder nitrogenase activity returned to pretreatment levels. Red alder root and nodule ADH activity was more than twice that of sitka alder under flooded conditions. Sitka alder showed extensive root mortality and leaf abscission over the same 20-day flooding period. Flooded red alder exhibited an initial decline in root RGR, but recovered between days 10 and 20 with the formation of adventitious roots. Furthermore, initiation of adventitious roots in red alder coincided with an increase in stomatal conductance without a similar recovery of carbon dioxide exchange rate. Sitka alder formed few adventitious roots, lost much of its root and leaf biomass, and showed no restoration of growth during flooding or recovery. Different responses of red and sitka alder to flooding serve as a partial explanation for the different patterns of distribution of these species and suggest some adaptations of red alder that permit flood tolerance.     

Growth responses of <Emphasis Type="Italic">Salix gracilistyla</Emphasis> cuttings to a range of substrate moisture and oxygen availability

The objective of this study is to determine the effects of substrate moisture and oxygen availability on growth traits of Salix gracilistyla Miquel, which colonizes gravel bars along rivers, the shoot growth schedule, biomass production, and resource allocation were examined under greenhouse conditions. We used four treatments representing a range of substrate moisture and oxygen availability: drought (D), flooding with standing water (FS), flooding with running water (FR), and control without drought or flooding (C). Cuttings in D stopped flushing and had low biomass production, reduced total leaf mass, and small leaves. Under anaerobic conditions, cuttings in FS stopped flushing and had low biomass production, small root biomass, low biomass allocation to roots, shallow roots, high biomass allocation to hypertrophied lenticels, and a few small, thick leaves. Under aerobic conditions, cuttings in FR showed continuous branch elongation and flushing, large biomass production, and large leaf biomass, similar to cuttings in C, in addition to low allocation to hypertrophied lenticels and many large leaves. The growth of cuttings was not inhibited by flooding of the roots throughout the experiment unless the conditions were anaerobic. Thus, cuttings respond to water stress under low moisture conditions by reducing the transpiration area and respond to flooding under low oxygen conditions by high allocation to hypertrophied lenticels and reduced transpiration area. Plasticity in the shoot growth schedule, biomass production, and resource allocation according to moisture conditions and the ability to develop hypertrophied lenticels upon flooding allow S. gracilistyla to colonize sites in which both desiccation and flooding occur.     

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     

Ethylene Production and Activation of Hydrolytic Enzymes during Acclimation of Maize Seedlings to Partial Flooding

The effects of partial flooding on the partial pressure of oxygen and carbon dioxide in water around the roots, ethylene production by intact maize (Zea mays L.) seedlings, the activities of hydrolytic enzymes (pectinase, xylanase, and cellulase) in adventitious roots, and the growth of adventitious and main roots were studied. Aggravated hypoxia resulted in the accelerated ethylene production and the activation of enzymes destroying cell walls in the adventitious roots; as a result, the latter changed their growth pattern. The conclusion is that the interrelated responses are adaptive ones, and the adventitious roots play a key role in plant adaptation.     

Roots of willow (Salix viminalis L.) show marked tolerance to oxygen shortage in flooded soils and in solution culture

Responses to soil flooding and oxygen shortage were studied in field, glasshouse and controlled environment conditions. Established stools ofSalix viminalis L., were compared at five field sites in close proximity but with contrasting water table levels and flooding intensities during the preceding winter. There was no marked effect of site on shoot extension rate, time to half maximum length or final length attained. When rooted cuttings were waterlogged for 4 weeks in a glasshouse, soil redox potentials quickly decreased to below zero. Shoot extension was slowed after a delay of 20 d, while, in the upper 100 mm of soil, formation and outgrowth of unbranched adventitious roots with enhanced aerenchyma development was promoted after 7 d. At depths of 100–200 mm and 200–300 mm, extension by existing root axes was halted by soil flooding, while adventitious roots from above failed to penetrate these deeper zones. After 4 weeks waterlogging, all arrested root tips recommenced elongation when the soil was drained; their extension rates exceeding those of roots that were well-drained throughout. Growth in fresh mass was also stimulated. The additional aerenchyma found in adventitious roots in the upper 100 mm of soil may have been ethylene regulated since gas space development was inhibited by silver nitrate, an ethylene action inhibitor. The effectiveness of aerenchyma was tested by blocking the entry of atmospheric oxygen into plants with lanolin applied to lenticels of woody shoots of plants grown in solution culture. Root extension was halved, while shoot growth remained unaffected. H Lambers Section editor     

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.