The aerenchymatous phellem of Lythrum salicaria (L.): a pathway for gas transport and its role in flood tolerance

While the importance of cortical aerenchyma in flood tolerance is well established, this pathway for gaseous exchange is often destroyed during secondary growth. For woody species, therefore, an additional pathway must develop for oxygen to reach submerged tissues. In this paper we examine the potential for the aerenchymatous phellem (cork) of Lythrum salicaria L. to provide a pathway for gas transport from shoots to roots and assess its importance in flood tolerance. Plants in which the continuity of the aerenchymatous phellem between shoots and roots was broken showed a significant reduction in oxygen levels in roots, but no difference in carbon dioxide levels compared with controls that retained an intact phellem. These plants also had a greater total shoot height and shoot dry weight, and an increase in shoot/root dry mass ratios compared with controls. Total dry weight was not significantly affected by this treatment. This study is the first to show that the aerenchymatous phellem can provide a pathway for gaseous exchange between roots and shoots and can influence plant morphology and patterns of resource allocation. This suggests that this tissue may play a significant role in the flood tolerance of a woody plant.     

Phenotypic plasticity in vegetative and reproductive traits in an invasive weed, Lythrum salicaria (Lythraceae), in response to soil moisture

In colonizing species, high phenotypic plasticity can contribute to survival and propagation in heterogenous adventive environments, and it has been suggested as a predictor of invasiveness. Observation of natural populations of an invasive species, Lythrum salicaria salicaria, indicated extensive variation in its growth and reproductive traits. Phenotypic plasticity of different life history traits of L. salicaria was investigated using vegetative clones of each of 12 genotypes from one population in Ontario, Canada. We chose soil moisture as the treatment factor because of its importance in wetland species and raised all 12 genotypes in each of four soil moisture treatments. We examined an array of vegetative and reproductive traits, including root and shoot mass, shoot and inflorescence length, total seed set, floral mass, and morphometric variables. All observed vegetative as well as reproductive traits demonstrated significant phenotypic plasticity in response to soil moisture treatment. Even the stigma-anther separation involved significant genotype by environment interactions, suggesting that soil moisture may modify the relative positions of anthers and stigma. Compared to vegetative traits, most reproductive traits demonstrated crossing reaction norms, implying that the average differences in those traits among genotypes vary with the environment maintaining the genetic variation in a population.     

Relationships among three pathways for resource acquisition and their contribution to plant performance in the emergent aquatic Plant Lythrum salicaria (L.)

Three pathways for resource acquisition exist in the emergent aquatic plant, Lythrum salicaria (L.); a subterranean root system, a free-floating adventitious root system, and arbuscular mycorrhiza (AM) fungal hyphae colonizing subterranean roots. This study examined the relationship(s) among these pathways and their contribution to plant performance. If the free-floating adventitious root system and/or AM fungi contribute to plant growth in wetland habitats, we predicted that their absence would result in a significant reduction in plant performance. Furthermore, if a reduction in resource uptake, effected by an absence of free-floating adventitious roots and/or AM fungi, is compensated for by increased allocation to remaining pathway(s) for resource uptake, we predicted altered patterns of resource allocation among shoots and the remaining pathway(s) for resource uptake. Contrary to our predications, plants experiencing adventitious root removal and/or grown in the absence of AM fungi generally had greater biomass and total shoot height than controls. Similarly, while levels of AM colonization and subterranean root biomass displayed a treatment effect, the observed responses did not correspond with our predictions. This was also true for shoot : subterranean root dry weight ratios. Our results indicate that there is interaction among the 3 pathways for resource acquisition in L. salicaria and an effect on plant performance. The adaptive significance of these characteristics is unclear, highlighting the potential difficulties in extrapolating from terrestrial to aquatic plant species and among aquatic plant species with potentially different life history strategies.     

The effects of flooding,plant traits,and predation on purple loosestrife leaf‐beetles

Classical biological weed control is based on the premise that introducing specialized natural enemies from the native range re‐establishes herbivore control of plant invaders, ultimately leading to negative population growth rates. Evidence from past biocontrol programs suggests that herbivores are not solely responsible for shaping plant demography. Diverse environmental conditions in the introduced range may not only affect demography, but also influence top‐down control of target plants. We investigated how flooding affects impacts of predators (top‐down) and plant quality (bottom‐up) on performance of two leaf‐beetles, Galerucella calmariensis L. and Galerucella pusilla Duftschmid (Coleoptera: Chrysomelidae: Galerucini), released in North America as biocontrol agents of purple loosestrife, Lythrum salicaria L. (Lythraceae). Predation and flooding regime have been linked to low leaf‐beetle recruitment at sites where insects failed to attain outbreak populations. Predator exclusion experiments at adjacent flooded and non‐flooded sites indicated a positive effect of flooding on leaf‐beetle survival for all developmental stages, whereas predator exposure had little effect. There was no difference in predation rates at sites with successful or failed purple loosestrife control, questioning the importance of predation in limiting growth and impact of these biocontrol agents’ populations. Effect of flooding on purple loosestrife quality was evaluated in a common garden study where plants were grown under different flooding treatments. Plants grown in flooded soil had higher water content and lower tannic acid concentration than plants grown in well‐drained soil. Consistent with field observations, leaf‐beetle oviposition rate and survival were higher on flooded plants. Results indicate that both bottom‐up and top‐down forces operate on Galerucella populations, yet their relative strength is mediated by flooding regime. Ignoring intricacies of plant‐herbivore and trophic interactions in the introduced range appears to be a major handicap for the improvement of weed biocontrol programs.     

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.     

ASSORTATIVE POLLINATION IN LYTHRUM

The pollination patterns of bees and butterflies were observed in an assemblage composed of 57 plants of Lythrum alatum and 30 plants of L. salicaria. Lythrum salicaria was preferred by both groups of pollinators as indicated by the greater number of flights from this species as compared to flights from L. alatum. The degree of assortative pollination was measured in both groups of pollinators and was greater in bees. Fidelity in bees appeared to be in response to species attractiveness, whereas in butterflies it appeared to be in response to species balance.     

Construction cost and invasive potential: comparing Lythrum salicaria (Lythraceae) with co-occurring native species along pond banks

Lythrum salicaria (purple loosestrife) is a nonindigenous invasive species characterized by prolific growth and abundance in marshy and riparian habitats across North America. Given its invasive success, we hypothesized this species may require less energy and/or use energy more efficiently for biomass construction than co-occurring noninvasive plant species. We measured leaf construction cost (CC), leaf mass per unit area (LMA), and leaf organic nitrogen and carbon content of L. salicaria and the five most abundant co-occurring species, Parthenocissus quinquefolia, Erigeron philadelphicus, Asclepias syriaca, Spiraea latifolia, and Solidago graminifolia, along dammed ponds in the Black Rock Forest, Cornwall, New York, USA. Lythrum salicaria, which was highly abundant (2.52 individuals/m(2)), exhibited significantly lower area-based leaf CC (44.47 ± 4.24 g glucose/m(2) leaf) than relatively less abundant species, suggesting energetics may influence its invasive success. Conversely, least abundant Solidago graminifolia (0.67 individuals/m(2)) exhibited the significantly highest leaf CC per unit leaf area (141.87 ± 39.21 g glucose/m(2) leaf). Overall, a negative correlation between species abundance and area-based leaf CC (r(2) = 0.73) indicated low energy requirements and/or high energy efficiency may influence relative abundance in the plant species studied. As it correlates with species abundance in this study, CC may be a useful tool for evaluating invasive potential.     

Evidence of hybridization between Lythrum salicaria (purple loosestrife) and L. alatum (winged loosestrife) in North America

BACKGROUND AND AIMS: Although Lythrum salicaria (purple loosestrife) was introduced to North America from Europe in the early 1800s, it did not become invasive until the 1930s. Whether hybridization with L. alatum (winged loosestrife) could have played a role in its ultimate spread was tested. METHODS: Six diagnostic morphological traits (flower number per axil, leaf placement, calyx pubescence, style type, plant height and leaf shape) were surveyed in 30 populations of Lythrum across eastern North America. Patterns of AFLP variation were also evaluated using five primer pairs in a 'global screen' of the same North American populations of L. salicaria and L. alatum described above, in L. salicaria from 11 European populations located in Germany, England, Ireland, Austria and Finland, and in six L. salicaria cultivars. KEY RESULTS: All of the North American L. salicaria populations had individuals with alternate leaf placement and 1-2 flowers per leaf axil, which have not been described in Eurasian L. salicaria but predominate in North American L. alatum. In addition, two L. salicaria populations were intermediate in height and leaf ratio between the typical L. salicaria and L. alatum populations in their native fields and when grown in a common greenhouse. In screens of variation patterns using 279 AFLPs, only two fragments were found that clearly supported introgression from L. alatum to L. salicaria. CONCLUSIONS: The evidence indicates that L. salicaria may have hybridized with L. alatum, but if so, only a small fraction of L. alatum genes have been retained in the genome of L. salicaria. This is unlikely to have led to a dramatic adaptive shift unless the introgression of a few key genes into L. salicaria stimulated a genomic reorganization. It is more likely that crossing among genotypes of L. salicaria from multiple introductions provided the necessary variability for new adaptations to arise.     

Increased relative abundance of an invasive competitor for pollination, Lythrum salicaria, reduces seed number in Mimulus ringens

When exotic plant species share pollinators with native species, competition for pollination may lower the reproductive success of natives by reducing the frequency and/or quality of visits they receive. Exotic species often become numerically dominant in plant communities, and the relative abundance of these potential competitors for pollination may be an important determinant of their effects on the pollination and reproductive success of co-occurring native species. Our study experimentally tests whether the presence and abundance of an invasive exotic, Lythrum salicaria L. (Lythraceae), influences reproductive success of a co-flowering native species, Mimulus ringens L. (Phrymaceae). We also examine the mechanisms of competition for pollination and how they may be altered by changes in competitor abundance. We found that the presence of Lythrum salicaria lowered mean seed number in Mimulus ringens fruits. This effect was most pronounced when the invasive competitor was highly abundant, decreasing the number of seeds per fruit by 40% in 2006 and 33% in 2007. Reductions in the number of seeds per fruit were likely due to reduced visit quality resulting from Mimulus pollen loss when bees foraged on neighboring Lythrum plants. This study suggests that visit quality to natives may be influenced by the presence and abundance of invasive flowering plants.     

Does flood tolerance explain tree species distribution in tropical seasonally flooded habitats?

In the tropics, seasonally flooded forests (SFF) harbor fewer tree species than terra firme (i.e. non-flooded) forests. The low species diversity of tropical flooded forests has been ascribed to the paucity of species with adaptations to tolerate flooding. To test the hypothesis that flooding is the only factor restricting most species from SFF, we compared plant morphological and physiological responses to flooding in 2-month old seedlings of 6 species common to SFF and 12 species common to terra firme forests. Although flooding impaired growth, total biomass, maximum root length and stomatal conductance in most species, responses varied greatly and were species-specific. For example, after 90 days, flooding reduced leaf area growth by 10-50% in all species, except in Tabebuia, a common species from non-flooded habitats. Similarly, flooding had a 5-45% negative effect on total biomass for all species, except in 1 SFF and 1 terra firme species both of which had more biomass under flooding. A principal component analysis, using the above responses to flooding, provided no evidence that SFF and terra firme species differed in their responses to flooding. Flooding also caused reductions in root growth for most species. Rooting depth and root: shoot ratios were significantly less affected by flooding in SFF than in terra firme species. Although flood tolerance is critical for survival in flooded habitats, we hypothesize that responses to post-flooding events such as drought might be equally important in seasonal habitats. Therefore, we suggest that the ability to grow roots under anoxia might be critical in predicting success in inundated habitats that also experience a strong dry season.     

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.     

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.     

FLOODWATER TEMPERATURE AND STEM LENTICEL HYPERTROPHY IN MANGIFERA INDICA (ANACARDIACEAE)

One-year-old seedling trees of Mangifera indica L. cv. Peach were exposed to floodwater temperatures of 15, 22.5, or 30 C for at least 13 days. Immediately prior to flooding, and at daily intervals thereafter, trees were visually examined for evidence of lenticel hypertrophy. Although lenticel hypertrophy was first apparent after 5 days of submergence at 30 C, and after 6 days at 22.5 C, the mean number of days of flooding until lenticel hypertrophy was first observed was 6.6 and 8.1 for the 30 and 22.5 C treatments, respectively. Even after 28 days, lenticel hypertrophy did not occur on plants flooded at 15 C. Initial stages of lenticel hypertrophy were characterized by development of intercellular spaces in the phellem and production of additional phellem tissue by increased phellogen activity. Later stages of hypertrophy were characterized by development of intercellular spaces in the phellem and cortex. Tree survival was not affected by floodwater temperature or lenticel hypertrophy.     

Interspecific pollinator movements reduce pollen deposition and seed production in Mimulus ringens (Phrymaceae)

Movement of pollinators between coflowering plant species may influence conspecific pollen deposition and seed set. Interspecific pollinator movements between native and showy invasive plants may be particularly detrimental to the pollination and reproductive success of native species. We explored the effects of invasive Lythrum salicaria on the reproductive success of Mimulus ringens, a wetland plant native to eastern North America. Pollinator flights between these species significantly reduced the amount of conspecific pollen deposited on Mimulus stigmas and the number of seeds in Mimulus fruits, suggesting that pollen loss is an important mechanism of competition for pollination. Although pollen loss is often attributed to pollen wastage on heterospecific floral structures, our novel findings suggest that grooming by bees as they forage on a competitor may also significantly reduce outcross pollen export and seed set in Mimulus ringens.     

Flooding effects on rapid responses of the invasive plant Alternanthera philoxeroides to defoliation

In experiments under controlled growth conditions it was examined how flooding affected the responses of the invasive plant Alternanthera philoxeroides to defoliation. In drained and flooded conditions, plants were subjected to five defoliation levels: 0, 10, 50, 90% removal of leaf tissue and apex removal (90% leaf tissue plus apical bud removal). Plants were harvested weekly for five weeks. In drained conditions, plant biomasses including total biomass, shoot biomass and root biomass after 50% defoliation rapidly recovered to the control plant level. They were significantly lower for the 90% defoliation and apex removal treatments compared to control plants throughout the experiment. In flooded conditions, total biomass and shoot biomass after 50% defoliation, 90% defoliation, and apex removal treatments could return to control plant levels before the end of the experiment. In 90% defoliation and apex removal treatments root to shoot biomass ratios of both drained and flooded plants were initially much higher than in control plants, but the difference disappeared rapidly. The final biomasses decreased with increased defoliation intensity in drained conditions, but no significant difference was generally found in any of the defoliation treatments in flooded conditions. The rapid re-growth of A. philoxeroides plants after defoliation may partly be responsible for its invasion success. However, defoliation capable of removing 90% of the leaf tissue may be desirable in restricting the growth of this invasive species in drained conditions.     

三峡库区河岸植物野古草(Arundinella anomala var. depauperata Keng)茎通气组织发生对水淹的响应

野古草(Arundinella anomala var. depauperata Keng)在三峡库区长江及其支流江(河)岸有广泛分布,对水淹有很好的耐受能力.有研究表明许多植物在水淹时通气组织发生增强,通气组织的产生改善了植株通气状况,提高了植物对水淹的抵御能力.为了研究水淹是否会影响野古草的通气组织发生以及野古草通气组织发生对水淹的反应,考察了不同水淹深度、不同水淹时间和不同水淹方式处理时野古草茎中通气组织的发生情况.实验中共设置3个水淹深度:不进行水淹(对照)、植株地下部分淹没、植株完全淹没于水下2m深处;5个淹没时间:植株被淹没的时间长度分别为5、10、20、30d和60d;2种水淹方式:连续水淹和间歇水淹.实验结果表明:(1)在无水淹情况下野古草茎中可以产生通气组织,通气组织产生随植株的生长而增强;水淹加快了野古草通气组织发生的进程,促进了野古草通气组织的提前发生.(2)野古草茎中通气组织并不会因为水淹的时间越长而产生越多,植株通气组织的大小达到一定程度后不再因水淹时间的增长而继续增大.(3)淹没深度对通气组织发生有一定影响,总的看来,地下部分淹没野古草植株的通气组织发生要强于完全淹没植株.(4)不同水淹方式对野古草通气组织发生的影响因水淹深度不同而有差异.在完全淹没情况下,连续水淹植株的通气组织比间歇水淹植株的通气组织发达;在地下部分淹没情况下,除水淹初期外,随水淹时间的延长,连续水淹植株通气组织发生与间歇水淹植株没有差异.     

Effects of soil fertility and flooding regime on the growth of Ambrosia trifida

Understanding the effects of abiotic environmental factors on invasive plants species traits is of importance for practical prevention. To examine the effects of soil fertility and flooding regime on the growth of Ambrosia trifida L., a mesocosm experiment was conducted for 18 weeks. Two levels of soil fertility (high and low) and three types of flooding regime (non-flooded, flooded, and periodically flooded) were prepared. Shoot height and dry weight of each plant were measured. We found both individual and interactive effects of soil fertility and flooding regime on the overall growth performance of A. trifida (p?<?0.05). The highest shoot height (154.7?±?4.4 cm) and total dry weight (TDW, 13.0?±?1.4 g) were obtained under high fertility and non-flooded condition. Height and weight were relatively low under flooding conditions (flooded and periodically flooded). In particular, shoot height (102.3?±?3.2 cm) and TDW (3.2?±?0.3 g) were the lowest under low fertility and periodically flooded condition. On the other hand, the ratio of above- to below-ground dry weight was relatively high under flooded conditions, showing the adaptive phenotypic plasticity. Adventitious root formation and more biomass allocation to shoots were a flooding-adaptive mechanism of A. trifida, well developed under high fertility condition. We suggest maintaining appropriate water regime and avoiding eutrophication in wetlands would be necessary to prevent A. trifida from invading. These findings will contribute to the conservation of biodiversity in wetlands by effective management of A. trifida.

     

Negative per capita effects of two invasive plants, Lythrum salicaria and Phalaris arundinacea, on the moth diversity of wetland communities

Invasive plants have been shown to negatively affect the diversity of plant communities. However, little is known about the effect of invasive plants on the diversity at other trophic levels. In this study, we examine the per capita effects of two invasive plants, purple loosestrife (Lythrum salicaria) and reed canary grass (Phalaris arundinacea), on moth diversity in wetland communities at 20 sites in the Pacific Northwest, USA. Prior studies document that increasing abundance of these two plant species decreases the diversity of plant communities. We predicted that this reduction in plant diversity would result in reduced herbivore diversity. Four measurements were used to quantify diversity: species richness (S), community evenness (J), Brillouin's index (H) and Simpson's index (D). We identified 162 plant species and 156 moth species across the 20 wetland sites. The number of moth species was positively correlated with the number of plant species. In addition, invasive plant abundance was negatively correlated with species richness of the moth community (linear relationship), and the effect was similar for both invasive plant species. However, no relationship was found between invasive plant abundance and the three other measures of moth diversity (J, H, D) which included moth abundance in their calculation. We conclude that species richness within, and among, trophic levels is adversely affected by these two invasive wetland plant species.     

Morphological changes and resource allocation of Zizania latifolia (Griseb.) Stapf in response to different submergence depth and duration

Plants around ponds, rivers and lakes are subjected to long-term partial or complete submergence. When they are flooded, water level affects the plants simultaneously with duration of submergence. Separate and interactive effects of water level and duration on the growth of the herbaceous perennial Zizania latifolia (Poaceae) were investigated by exposing the plants in greenhouse water tanks to submergence in different water depths and for different time-spans. The plants exhibited great shoot elongation upon submergence and prolonged flood duration, and the basal tiller number of the species decreased with higher water levels. Submergence treatment advanced the flowering date and increased the inflorescence number. Plant total biomass did not differ among all the treatments, while the root:shoot ratio decreased with increased water level, prolonged duration of submergence and their interaction. The high plasticity in morphology and shifts in reproductive strategy and biomass allocation enabled the Zizania plants to survive the compound effect of flooding height and duration. This may explain the occurrence of this species in habitats subjected to long-term flooding. The results obtained in this experiment will contribute to understanding the impact of flooding dynamics on plants and the ways of adaptation responses to prolonged waterlogging.     

Effects of soil moisture regimes on photosynthesis and growth in cattail (Typha latifolia)

Both waterlogging and water deficiency are major environmental factors affecting plant growth and functioning in many wetland and floodplain ecosystems across North America. Wetland plants possess various characteristics that enable them to survive and function in the intermittently flooded wetland environments, while their sensitivity to drought has received less attention. The present study quantified the photosynthetic and growth responses of cattail (Typha latifolia), an important species of freshwater wetlands, to a wide range of soil moisture regimes. In addition, changes in the efficiency of photosynthetic apparatus following initiation of the treatments were investigated. Under greenhouse conditions, seedlings were subjected to four soil moisture regimes: (1) drained (control), (2) continuous flooding, (3) periodic flooding, and (4) periodic drought. Results indicated that dark fluorescence yield was increased in response to periodic drought, while it showed decreases under continuous flooding. Net photosynthesis and stomatal conductance were enhanced by continuous flooding and periodic flooding. In contrast, these parameters exhibited reduction under periodic drought. In addition, leaf chlorophyll content was adversely affected by periodic drought. Recovery of net photosynthesis was noted, along with enhanced height growth, in both continuously and periodically flooded plants. Meanwhile, continuous flooding enhanced biomass production while periodic drought led to biomass reduction. Periodic drought also contributed to substantial reduction in root growth compared with shoot growth. Therefore, the combined photosynthetic performance and growth responses of cattail are likely to contribute to the ability of this species to thrive in flooded condition but be susceptive to periodic drought.