Effects of different humic substances concentrations on root anatomy and Cd accumulation in seedlings of Avicennia germinans (black mangrove).

Mangrove areas are among most threatened tropical ecosystems worldwide. Among polluting agents Cadmium is often found in high concentrations in mangrove sediments. Humic substances, complex biomolecules formed in soil and sediments during animal and plant residuals decomposition, have a known biostimulant activity and can be adopted to counteract various plant stresses. This study explores, in controlled conditions, the effect of humic substances on Avicennia germinans seedlings, with or without cadmium contamination. Humic compounds significantly changed plant root architecture, and, when coupled with cadmium, root anatomy and Cortex to Vascular Cylinder diameter ratio. These modifications led to lower Cd uptake by humic substances-treated plants. Humic substances amendment could be effective, depending on their concentrations, on improving plant health in mangrove areas, for forest recuperation and/or dredged sediments phytoremediation purposes.

Leaf miner-induced morphological, physiological and molecular changes in mangrove plant Avicennia marina (Forsk.) Vierh.

Avicennia marina (Forsk.) Vierh is a widespread mangrove species along the southeast coasts of China. Recently, the outbreak of herbivorous insect, Phyllocnistis citrella Stainton, a leaf miner, have impacted on the growth of A. marina. Little is reported about the responses of A. marina to leaf miner infection at the biochemical, physiological and molecular levels. Here, we reported the responses of A. marina to leaf miner infection from the aspects of leaf structure, photosynthesis, and antioxidant system and miner responsive genes expression. A. marina leaves attacked by the leaf miner exhibited significant decreases in chlorophyll, carbon and nitrogen contents, as well as a decreased photosynthetic rate. Scanning and transmission electron microscopic observations revealed that the leaf miner only invaded the upper epidermis and destroyed the epidermal cell, which lead to the exposure of salt glands. In addition, the chloroplasts of mined leaves (ML) were swollen and the thylakoids degraded. The maximal net photosynthetic rate, stomatal conductance (Gs), carboxylation efficiency (CE), dark respiration (Rd), light respiration (Rp) and quantum yields (AQE) significantly decreased in the ML, whereas the light saturation point (Lsp), light compensation point (Lcp), water loss and CO2 compensation point (Г) increased in the ML. Moreover, chlorophyll fluorescence features also had been changed by leaf miner attacks. Interestingly, higher generation rate of O2ˉ· and lower antioxidant enzyme expression in the mined portion (MP) were found; on the contrary, higher H2O2 level and higher antioxidant enzyme expression in the non-mined portion (NMP) were revealed, implying that the NMP may be able to sense that the leaf miner attacks had happened in the MP of the A. marina leaf via H2O2 signaling. Besides, the protein expression of glutathione S-transferase (GST) and the glutathione (GSH) content were increased in the ML. In addition, insect resistance-related gene expression such as chitinase 3, RAR1, topless and PIF3 had significantly increased in the ML. Taken together, our data suggest that leaf miners could significantly affect leaf structure, photosynthesis, the antioxidant system and miner responsive gene expression in A. marina leaves.

Plumbing the depths: extracellular water storage in specialized leaf structures and its functional expression in a three-domain pressure -volume relationship.

A three-domain pressure-volume relationship (PV curve) was studied in relation to leaf anatomical structure during dehydration in the grey mangrove, Avicennia marina. In domain 1, relative water content (RWC) declined 13% with 0.85 MPa decrease in leaf water potential, reflecting a decrease in extracellular water stored primarily in trichomes and petiolar cisternae. In domain 2, RWC decreased by another 12% with a further reduction in leaf water potential to -5.1 MPa, the turgor loss point. Given the osmotic potential at full turgor (-4.2 MPa) and the effective modulus of elasticity (~40 MPa), domain 2 emphasized the role of cell wall elasticity in conserving cellular hydration during leaf water loss. Domain 3 was dominated by osmotic effects and characterized by plasmolysis in most tissues and cell types without cell wall collapse. Extracellular and cellular water storage could support an evaporation rate of 1 mmol m-2 s-1 for up to 54 and 50 min, respectively, before turgor loss was reached. This study emphasized the importance of leaf anatomy for the interpretation of PV curves, and identified extracellular water storage sites that enable transient water use without substantive turgor loss when other factors, such as high soil salinity, constrain rates of water transport.

Proteomic Characterisation of the Salt Gland-Enriched Tissues of the Mangrove Tree Species Avicennia officinalis.

Plant salt glands are nature's desalination devices that harbour potentially useful information pertaining to salt and water transport during secretion. As part of the program toward deciphering secretion mechanisms in salt glands, we used shotgun proteomics to compare the protein profiles of salt gland-enriched (isolated epidermal peels) and salt gland-deprived (mesophyll) tissues of the mangrove species Avicennia officinalis. The purpose of the work is to identify proteins that are present in the salt gland-enriched tissues. An average of 2189 and 977 proteins were identified from the epidermal peel and mesophyll tissues, respectively. Among these, 2188 proteins were identified in salt gland-enriched tissues and a total of 1032 selected proteins were categorized by Gene Ontology (GO) analysis. This paper reports for the first time the proteomic analysis of salt gland-enriched tissues of a mangrove tree species. Candidate proteins that may play a role in the desalination process of the mangrove salt glands and their potential localization were identified. Information obtained from this study paves the way for future proteomic research aiming at elucidating the molecular mechanism underlying secretion in plant salt glands. The data have been deposited to the ProteomeXchange with identifier PXD000771.

Root biomechanics in Rhizophora mangle: anatomy, morphology and ecology of mangrove's flying buttresses.

BACKGROUND AND AIMS: Rhizophora species of mangroves have a conspicuous system of stilt-like roots (rhizophores) that grow from the main stem and resemble flying buttresses. As such, the development of rhizophores can be predicted to be important for the effective transmission of dynamic loads from the top of the tree to the ground, especially where the substrate is unstable, as is often the case in the habitats where Rhizophora species typically grow. This study tests the hypothesis that rhizophore architecture in R. mangle co-varies with their proximity to the main stem, and with stem size and crown position. METHODS: The allometry and wood mechanical properties of R. mangle (red mangrove) trees growing in a mangrove basin forest within a coastal lagoon in Mexico were compared with those of coexisting, non-buttressed mangrove trees of Avicennia germinans. The anatomy of rhizophores was related to mechanical stress due to crown orientation (static load) and to prevailing winds (dynamic load) at the study site. KEY RESULTS: Rhizophores buttressed between 10 and 33 % of tree height. There were significant and direct scaling relationships between the number, height and length of rhizophores vs. basal area, tree height and crown area. Wood mechanical resistance was significantly higher in the buttressed R. mangle (modulus of elasticity, MOE = 18·1 ± 2 GPa) than in A. germinans (MOE = 12·1 ± 0·5 GPa). Slenderness ratios (total height/stem diameter) were higher in R. mangle, but there were no interspecies differences in critical buckling height. When in proximity to the main stem, rhizophores had a lower length/height ratio, higher eccentricity and higher xylem/bark and pith proportions. However, there were no directional trends with regard to prevailing winds or tree leaning. CONCLUSIONS: In comparison with A. germinans, a tree species with wide girth and flare at the base, R. mangle supports a thinner stem of higher mechanical resistance that is stabilized by rhizophores resembling flying buttresses. This provides a unique strategy to increase tree slenderness and height in the typically unstable substrate on which the trees grow, at a site that is subject to frequent storms.

Growth responses of the mangrove Avicennia marina to salinity: development and function of shoot hydraulic systems require saline conditions.

BACKGROUND AND AIMS: Halophytic eudicots are characterized by enhanced growth under saline conditions. This study combines physiological and anatomical analyses to identify processes underlying growth responses of the mangrove Avicennia marina to salinities ranging from fresh- to seawater conditions. METHODS: Following pre-exhaustion of cotyledonary reserves under optimal conditions (i.e. 50% seawater), seedlings of A. marina were grown hydroponically in dilutions of seawater amended with nutrients. Whole-plant growth characteristics were analysed in relation to dry mass accumulation and its allocation to different plant parts. Gas exchange characteristics and stable carbon isotopic composition of leaves were measured to evaluate water use in relation to carbon gain. Stem and leaf hydraulic anatomy were measured in relation to plant water use and growth. KEY RESULTS: Avicennia marina seedlings failed to grow in 0-5% seawater, whereas maximal growth occurred in 50-75% seawater. Relative growth rates were affected by changes in leaf area ratio (LAR) and net assimilation rate (NAR) along the salinity gradient, with NAR generally being more important. Gas exchange characteristics followed the same trends as plant growth, with assimilation rates and stomatal conductance being greatest in leaves grown in 50-75% seawater. However, water use efficiency was maintained nearly constant across all salinities, consistent with carbon isotopic signatures. Anatomical studies revealed variation in rates of development and composition of hydraulic tissues that were consistent with salinity-dependent patterns in water use and growth, including a structural explanation for low stomatal conductance and growth under low salinity. CONCLUSIONS: The results identified stem and leaf transport systems as central to understanding the integrated growth responses to variation in salinity from fresh- to seawater conditions. Avicennia marina was revealed as an obligate halophyte, requiring saline conditions for development of the transport systems needed to sustain water use and carbon gain.

Modelling reveals endogenous osmotic adaptation of storage tissue water potential as an important driver determining different stem diameter variation patterns in the mangrove species Avicennia marina and Rhizophora stylosa.

BACKGROUND: Stem diameter variations are mainly determined by the radial water transport between xylem and storage tissues. This radial transport results from the water potential difference between these tissues, which is influenced by both hydraulic and carbon related processes. Measurements have shown that when subjected to the same environmental conditions, the co-occurring mangrove species Avicennia marina and Rhizophora stylosa unexpectedly show a totally different pattern in daily stem diameter variation. METHODS: Using in situ measurements of stem diameter variation, stem water potential and sap flow, a mechanistic flow and storage model based on the cohesion-tension theory was applied to assess the differences in osmotic storage water potential between Avicennia marina and Rhizophora stylosa. KEY RESULTS: Both species, subjected to the same environmental conditions, showed a resembling daily pattern in simulated osmotic storage water potential. However, the osmotic storage water potential of R. stylosa started to decrease slightly after that of A. marina in the morning and increased again slightly later in the evening. This small shift in osmotic storage water potential likely underlaid the marked differences in daily stem diameter variation pattern between the two species. CONCLUSIONS: The results show that in addition to environmental dynamics, endogenous changes in the osmotic storage water potential must be taken into account in order to accurately predict stem diameter variations, and hence growth.

How to catch the patch? A dendrometer study of the radial increment through successive cambia in the mangrove Avicennia.

BACKGROUND AND AIMS: Successive vascular cambia are involved in the secondary growth of at least 200 woody species from >30 plant families. In the mangrove Avicennia these successive cambia are organized in patches, creating stems with non-concentric xylem tissue surrounded by internal phloem tissue. Little is known about radial growth and tree stem dynamics in trees with this type of anatomy. This study aims to (1) clarify the process of secondary growth of Avicennia trees by studying its patchiness; and (2) study the radial increment of Avicennia stems, both temporary and permanent, in relation to local climatic and environmental conditions. A test is made of the hypothesis that patchy radial growth and stem dynamics enable Avicennia trees to better survive conditions of extreme physiological drought. Methods Stem variations were monitored by automatic point dendrometers at four different positions around and along the stem of two Avicennia marina trees in the mangrove forest of Gazi Bay (Kenya) during 1 year. KEY RESULTS: Patchiness was found in the radial growth and shrinkage and swelling patterns of Avicennia stems. It was, however, potentially rather than systematically present, i.e. stems reacted either concentrically or patchily to environment triggers, and it was fresh water availability and not tidal inundation that affected radial increment. CONCLUSIONS: It is concluded that the ability to develop successive cambia in a patchy way enables Avicennia trees to adapt to changes in the prevailing environmental conditions, enhancing its survival in the highly dynamic mangrove environment. Limited water could be used in a more directive way, investing all the attainable resources in only some locations of the tree stem so that at least at these locations there is enough water to, for example, overcome vessel embolisms or create new cells. As these locations change with time, the overall functioning of the tree can be maintained.

Successive cambia: a developmental oddity or an adaptive structure?

BACKGROUND: Secondary growth by successive cambia is a rare phenomenon in woody plant species. Only few plant species, within different phylogenetic clades, have secondary growth by more than one vascular cambium. Often, these successive cambia are organised concentrically. In the mangrove genus Avicennia however, the successive cambia seem to have a more complex organisation. This study aimed (i) at understanding the development of successive cambia by giving a three-dimensional description of the hydraulic architecture of Avicennia and (ii) at unveiling the possible adaptive nature of growth by successive cambia through a study of the ecological distribution of plant species with concentric internal phloem. RESULTS: Avicennia had a complex network of non-cylindrical wood patches, the complexity of which increased with more stressful ecological conditions. As internal phloem has been suggested to play a role in water storage and embolism repair, the spatial organisation of Avicennia wood could provide advantages in the ecologically stressful conditions species of this mangrove genus are growing in. Furthermore, we could observe that 84.9% of the woody shrub and tree species with concentric internal phloem occurred in either dry or saline environments strengthening the hypothesis that successive cambia provide the necessary advantages for survival in harsh environmental conditions. CONCLUSIONS: Successive cambia are an ecologically important characteristic, which seems strongly related with water-limited environments.

Effect of wastewater discharge on root anatomy and radial oxygen loss (ROL) patterns of three mangrove species in southern China.

The effects of wastewater discharge on radial oxygen loss (ROL) and root anatomy varied among mangrove species. ROL of Bruguiera gymnorrhiza (L) increased from 22.44 ng cm(-2) min(-1) in the control (just fresh water) to 31.09 ng cm(-2) min(-1) when received normal wastewater (NW) and to 44.22 ng cm(-2) min(-1) when treated with strong wastewater (10NW). However, discharge of both NW and 10NW caused 28% decreases of ROL in the root tip of Excoecaria agallocha L., and the decreases in Acanthus ilicifolius L were even more significant, with 45% when treated by 10NW The changes of ROL were related to the root anatomy. Among three species, A. ilicifolius had the highest proportional cross-sectional area of aerenchyma air spaces, suggesting that the internal oxygen transfer to root tip was the fastest. However, the area of aerenchyma air spaces in the root tip of 10NW treated A. ilicifolius was significantly reduced while area of epidermis and hypodermis (E + H) increased leading to less oxygen supply to root tip. Compared to B. gymnorrhiza and E. agallocha, the (E + H) layer of A. ilicifolius was the thinnest, and the cells without suberized walls were loosely packed in all three treatments. These results suggested that the root anatomy and ROL of B. gymnorrhiza was least affected by wastewater discharge, followed by E. agallocha, and A. ilicifolius was the most susceptible species thus was not suitable for treating strong wastewater.

[Costa Rica mangroves: the north Pacific]. / Los manglares de Costa Rica: el Pacífico norte.

Costa Rica has mangrove forests on both the Caribbean and Pacific coasts. The Pacific side has 99% of the mangrove area of the country. In this review we compile available information on the mangroves of the north Pacific coast of Costa Rica, from Bahía Salinas, on the border with Nicaragua, to the tip of the Peninsula de Nicoya at Cabo Blanco. We provide information on the location of the mangroves and all available information for each mangrove forest. These mangrove communities are smaller in extension and tree sizes, and have lower diversity compared to the mangroves on the southern section of the Pacific coast of Costa Rica. The dominant species are Rhizophora mangle and Rhizophora racemosa along the canal edges, backed by Avicennia germinans, and farther inland Avicennia bicolor, Laguncularia racemosa and Conocarpus erectus. At Potrero Grande a healthy population of Pelliciera rhizophorae, a rare species, has been reported. We recognized 38 mangrove communities in this part of the country, based on the National Wetland Inventory, published papers, field observations, theses, technical reports, and the national topographic maps (1:50,000, Instituto Geográfico Nacional). Relatively detailed information could be found for only five mangrove forests, for 14 more only prelimary and incomplete lists of plants and in some cases of animal species are available, for nine there is even less information, and for nine more only their location is known, which in some cases was not correct. Detail mapping, characterization of the vegetation and fauna, physiological studies, analyses of biogeochemical and physical processes, economic valuations, and determination of the health status of the mangrove of the northern Pacific coast, as well as for the rest of Costa Rica, are neccesary and urgent.

Los manglares de Costa Rica: el Pacífico norte

Abstract: Costa Rica mangroves: the north Pacific. Costa Rica has mangrove forests on both the Caribbean and Pacific coasts. The Pacific side has 99% of the mangrove area of the country. In this review we compile available information on the mangroves of the north Pacific coast of Costa Rica, from Bahía Salinas, on the border with Nicaragua, to the tip of the Peninsula de Nicoya at Cabo Blanco. We provide information on the location of the mangroves and all available information for each mangrove forest. These mangrove communities are smaller in extension and tree sizes, and have lower diversity compared to the mangroves on the southern section of the Pacific coast of Costa Rica. The dominant species are Rhizophora mangle and Rhizophora racemosa along the canal edges, backed by Avicennia germinans, and farther inland Avicennia bicolor, Laguncularia racemosa and Conocarpus erectus. At Potrero Grande a healthy population of Pelliciera rhizophorae, a rare species, has been reported. We recognized 38 mangrove communities in this part of the country, based on the National Wetland Inventory, published papers, field observations, theses, technical reports, and the national topographic maps (1:50 000, Instituto Geográfico Nacional). Relatively detailed information could be found for only five mangrove forests, for 14 more only prelimary and incomplete lists of plants and in some cases of animal species are available, for nine there is even less information, and for nine more only their location is known, which in some cases was not correct. Detail mapping, characterization of the vegetation and fauna, physiological studies, analyses of biogeochemical and physical processes, economic valuations, and determination of the health status of the mangrove of the northern Pacific coast, as well as for the rest of Costa Rica, are neccesary and urgent. Rev. Biol. Trop. 57 (3): 473-488. Epub 2009 September 30.

Costa Rica tiene bosques de manglar en las costas Caribe y Pacífica. El 99% de los manglares del país se encuentran en la costa Pacífica. En este trabajo recopilamos la información disponible sobre los manglares del Pacífico norte de Costa Rica, desde la frontera con Nicaragua hasta la Reserva Absoluta de Cabo Blanco, en la punta sur de la Península de Nicoya. La ubicación de los manglares y cualquier información disponible para cada manglar es resumida. El clima de esta región es seco con estaciones de lluvia y sequía muy bien definidas. Los manglares del norte son relativamente pequeños en extensión y tamaño de árboles, y de baja diversidad comparados con los del Pacífico sur de Costa Rica. Las principales especies de mangle son: Rhizophora mangle, Rhizophora racemosa a lo largo de los bordes de los canales, seguido por Avicennia germinans, y más tierra adentro Avicennia bicolor, Laguncularia racemosa y Conocarpus erectus. En el manglar de Potrero Grande se informa de una población saludable de la especie de Pelliciera rhizophorae, especies muy poco abundante. Reconocemos 38 comunidades de manglares en el Pacífico norte de Costa Rica basado en el Inventario Nacional de Humedales, artículos publicados, observaciones de campo, tesis, informes técnicos y los mapas 1: 50 000 del Instituto Geográfico Nacional. Se pudo encontrar información relativamente detallada pero fragmentada de únicamente cinco manglares (los descritos en esta publicación), de 14 solamente hay listas preliminares e incompletas de plantas y en algunos casos de animales, de nueve manglares más hay información todavía más limitada, y de otros nueve se conoce solamente su ubicación, que en algunos casos se comprobó que era incorrecta. Trabajos de mapeo detallado, caracterización de la vegetación y la fauna, estudios fisiológicos, análisis de procesos biogeoquímicos, evaluaciones económicas, y determinación del estado de salud de los manglares del Pacífico norte de Costa Rica, y del resto del país son necesarios y urgentes.

A patchy growth via successive and simultaneous cambia: key to success of the most widespread mangrove species Avicennia marina?

BACKGROUND AND AIMS: Secondary growth via successive cambia has been intriguing researchers for decades. Insight into the mechanism of growth layer formation is, however, limited to the cellular level. The present study aims to clarify secondary growth via successive cambia in the mangrove species Avicennia marina on a macroscopic level, addressing the formation of the growth layer network as a whole. In addition, previously suggested effects of salinity on growth layer formation were reconsidered. METHODS: A 1-year cambial marking experiment was performed on 80 trees from eight sites in two mangrove forests in Kenya. Environmental (soil water salinity and nutrients, soil texture, inundation frequency) and tree characteristics (diameter, height, leaf area index) were recorded for each site. Both groups of variables were analysed in relation to annual number of growth layers, annual radial increment and average growth layer width of stem discs. KEY RESULTS: Between trees of the same site, the number of growth layers formed during the 1-year study period varied from only part of a growth layer up to four growth layers, and was highly correlated to the corresponding radial increment (0-5 mm year(-1)), even along the different sides of asymmetric stem discs. The radial increment was unrelated to salinity, but the growth layer width decreased with increasing salinity and decreasing tree height. CONCLUSIONS: A patchy growth mechanism was proposed, with an optimal growth at distinct moments in time at different positions around the stem circumference. This strategy creates the opportunity to form several growth layers simultaneously, as observed in 14 % of the studied trees, which may optimize tree growth under favourable conditions. Strong evidence was provided for a mainly endogenous trigger controlling cambium differentiation, with an additional influence of current environmental conditions in a trade-off between hydraulic efficiency and mechanical stability.

The role of freezing in setting the latitudinal limits of mangrove forests.

Mangrove trees dominate coastal vegetation in tropical regions, but are completely replaced by herbaceous salt marshes at latitudes above 32 degrees N and 40 degrees S. Because water deficit can increase damage caused by freezing, we hypothesized that mangroves, which experience large deficits as a result of saline substrates, would suffer freeze-induced xylem failure. Vulnerability to freeze-induced xylem embolism was examined in the most poleward mangrove species in North America, in an area where freezing is rare but severe, and in Australia, in an area where freezing is frequent but mild. Percentage loss in hydraulic conductivity was measured following manipulations of xylem tension; xylem sap ion concentration was determined using X-ray microanalysis. Species with wider vessels suffered 60-100% loss of hydraulic conductivity after freezing and thawing under tension, while species with narrower vessels lost as little as 13-40% of conductivity. These results indicate that freeze-induced embolism may play a role in setting the latitudinal limits of distribution in mangroves, either through massive embolism following freezing, or through constraints on water transport as a result of vessel size.

The influence of radical architecture on cadmium bioaccumulation in the black mangrove, Avicennia germinans L.

Two groups of Avicennia germinans plants with differences in the radical architecture were exposed under hydroponic conditions to 95ppm of cadmium (Cd) for a period of 24h. Later, Cd concentration in roots, stems and leaves was determined by graphite furnace atomic absorption spectrophotometry. Our results showed that, for both groups of plants, the roots accumulated higher concentration of Cd as compared to stems and leaves, though, the plants of group B displayed enhanced radical architecture, better growth performance, and lower Cd concentration as compared to plants of group A. In contrast, low values of leaves/roots Cd transportation index, and bioaccumulation factor were found in plants of group B. These results suggest that the higher radical architecture developed in plants of group B might better adjust the uptake of Cd as a result of an integrated network of multiple response processes for instances, production of organic acids, antioxidative replay, cell-wall lignification and/or suberization. Further studies will be focused in understanding the role of the radical system in mangrove plants with the rhizosphere activation and root adsorption to soil Cd under natural conditions.

Leaf gas exchange characteristics of three neotropical mangrove species in response to varying hydroperiod.

We determined how different hydroperiods affected leaf gas exchange characteristics of greenhouse-grown seedlings (2002) and saplings (2003) of the mangrove species Avicennia germinans (L.) Stearn., Laguncularia racemosa (L.) Gaertn. f., and Rhizophora mangle L. Hydroperiod treatments included no flooding (unflooded), intermittent flooding (intermittent), and permanent flooding (flooded). Plants in the intermittent treatment were measured under both flooded and drained states and compared separately. In the greenhouse study, plants of all species maintained different leaf areas in the contrasting hydroperiods during both years. Assimilation-light response curves indicated that the different hydroperiods had little effect on leaf gas exchange characteristics in either seedlings or saplings. However, short-term intermittent flooding for between 6 and 22 days caused a 20% reduction in maximum leaf-level carbon assimilation rate, a 51% lower light requirement to attain 50% of maximum assimilation, and a 38% higher demand from dark respiration. Although interspecific differences were evident for nearly all measured parameters in both years, there was little consistency in ranking of the interspecific responses. Species by hydroperiod interactions were significant only for sapling leaf area. In a field study, R. mangle saplings along the Shark River in the Everglades National Park either demonstrated no significant effect or slight enhancement of carbon assimilation and water-use efficiency while flooded. We obtained little evidence that contrasting hydroperiods affect leaf gas exchange characteristics of mangrove seedlings or saplings over long time intervals; however, intermittent flooding may cause short-term depressions in leaf gas exchange. The resilience of mangrove systems to flooding, as demonstrated in the permanently flooded treatments, will likely promote photosynthetic and morphological adjustment to slight hydroperiod shifts in many settings.