Effects of stream flooding on the distribution and diversity of groundwater‐dependent vegetation in riparian areas

1. Effects of the frequency and duration of flooding on the structural and functional characteristics of riparian vegetation were studied at four sites (n = 80, 50 × 50 cm, plots) along medium‐sized naturally meandering lowland streams. Special focus was on rich fens, which – due to their high species richness – are of high priority in nature conservation. 2. Reed beds, rich fens and meadows were all regularly flooded during the 20‐year study period, with a higher frequency in reed bed areas than in rich fen and meadow areas. In rich fens, species richness was higher in low frequency flooded areas (≤3 year^?1) than in areas with a high frequency of flooding (>3 year^?1) or no flooding, whereas species richness in reed beds and meadows was unaffected by flood frequency. 3. The percentage of stress‐tolerant species was higher in low intensity flooded rich fen areas than in high intensity and non‐flooded areas, indicating that the higher species richness in low frequency flooded rich fens was caused by competitive release. We found no indication that increased productivity was associated with high flooding frequencies. 4. We conclude that the restoration of morphological features in stream channels to increase the flooding regime can be beneficial for protected vegetation within riparian areas, but also that groundwater discharge thresholds and critical levels for protected vegetation should be identified and considered when introducing stream ecosystem restoration plans.     

Impact of flooding on potential and realised grassland species richness

In order to reduce flood risk, river management policies advise floodplain restoration and the recreation of water retention areas. These measures may also offer opportunities for the restoration of species-rich floodplain habitats through rewetting and the restoration of flood dynamics. The potential to enhance biodiversity in such flood restoration areas is, however, still subject to debate. In this paper we investigate whether flooding along a small altered lowland river can contribute to the potential and realised species richness of semi-natural meadows. We compare the seed bank and vegetation composition of flooded and non-flooded semi-natural meadows and test the hypothesis that flooding contributes to an input of diaspores into the meadow seed banks, thereby promoting seed density and potential species richness. Furthermore we hypothesise that, where habitat conditions are suitable, flooding leads to a higher realised species richness. Results showed that seed densities in flooded meadows were significantly higher than in non-flooded meadows. The seed banks of flooded meadows also contained a higher proportion of exclusively hydrochorous species. However, the seed bank species richness, as well as the species richness realised in the vegetation did not differ significantly between flooded and non-flooded meadows. Finally, the seed bank and standing vegetation of flooded sites showed larger differences in species composition and Ellenberg nitrogen distribution than non-flooded sites. From these results we conclude that, although flooding does contribute to the density and composition of the seed bank, most imported seeds belong to only a few species. Therefore, it is unlikely that flooding substantially enhances the potential species richness. Furthermore, even if new species are imported as seeds into the seed bank, it seems unlikely that they would be able to establish in the standing vegetation. However, it is unclear which factors impede the establishment of imported species in the vegetation. The implications of our findings for flood meadow restoration are discussed.     

Some physiological and morphological responses of Pyrus boissieriana to flooding

European pear is a flooding-sensitive species, and for its cultivation in lowland areas, it is necessary to carry out the grafting of scions of commercial pear varieties into rootstocks belonging to flooding-tolerant wild pear species. Flooding tolerance of Pyrus boissieriana—a type of wild pear—was studied as a promissory rootstock for commercial pear. For this purpose, 3-month-old plants of P. boissieriana were subjected for 30 days to control (C), well-irrigated treatment, short-term (15 days) flooding plus 15 days recovery (F + R) and long-term (30 days) continuous flooding (F). Physiological performance, plant morphological changes and biomass accumulation were assessed. Results showed that, although stomatal conductance, transpiration and photosynthesis were progressively decreased by flooding, when flooding was short term (i.e., 2 weeks, F + R treatment) plants were able to adequately recover their physiological activity (50–74 % with respect to controls). In contrast, when plants continued to be flooded (F treatment), the physiological activity became null and the plants died quickly after the water subsided. Adventitious rooting was the most conspicuous registered morphological response to flooding, despite that flooded plants had shorter shoots and roots than control plants. Leaf and root biomass were 63 and 89 % higher under short-term flooding (F + R) than under continuous flooding (F), condition in which plants did not survive. In conclusion, P. boissieriana appears to be a promising species for its use as rootstock of commercial pear in lowland areas prone to flooding of up to 2 weeks. However, if the flooding period is extended, plants of this species are at risk of perishing.     

Flooding tolerance of Tabebuia cassinoides: Metabolic, morphological and growth responses

Tabebuia cassinoides (Lam.) DC (Bignoniaceae) is a tree species that occurs in swampy areas of the coastal “restinga” in SE Brazil (a coastal sandy plains scrub and forest formation). To elucidate possible adaptive strategies that enable this species to occupy areas subjected to seasonal or perennial waterlogging, metabolic, morphological and growth responses of plants under flooding conditions were studied. The root system of T. cassinoides plants presented elevated amounts of ethanol (10.6 μmol g⁻¹ fresh wt) only in the first 5 d of soil water saturation. The two-fold increase in ethanol production under flooding was corroborated by an increase in ADH activity in the same period. Lactic acid concentrations did not change significantly during four months of flooding treatment. The decrease of alcoholic fermentation under hypoxia was associated with the appearing of new roots. The induction of aerenchyma formation in roots developed under flooding conditions, allowed oxygen transport from the shoot to these organs, thus maintaining an aerobic respiration. We conclude that this characteristic and the capacity to oxidize the rhizosphere are probably responsible for the survival and growth of plants while flooded and for their success in an environment, which restricts the presence of the majority of competing tree species.     

Juxtaposition and Disturbance: Disentangling the Determinants of Lizard Community Structure

Disturbance alters the structure and dynamics of communities. Here, we examined the effects of seasonal flooding on the lizard community structure by comparing two adjacent habitats, a seasonally flooded and a non‐flooded forest, in a Cerrado–Amazon ecotone area, the Cantão State Park, Tocantins state, Brazil. Despite the strong potential impact of seasonal flooding, the only significant environmental difference detected was more termite mounds in non‐flooded forests. Species richness was significantly higher in the non‐flooded forest. Colobosaura modesta, followed by Mabuya frenata and Anolis brasiliensis, were the only species that differed in number of captures between sites. Colobosaura modesta was exclusively found in the non‐flooded forest, while Anolis brasiliensis was the most captured in the flooded forest. Mabuya frenata is indicated as an indicator species in the flooded forest, and Colobosaura modesta in the non‐flooded forest. We found a significant association between lizard abundances and habitat characteristics, with flooding, canopy cover, and logs being the best predictors. A phylogenetic community structure analysis indicated a lack of structure in both lizard assemblages. Overall, we show that seasonal flooding can strongly impact species richness and species occurrence patterns, but not phylogenetic community structure. The Amazon–Cerrado transition is undergoing pronounced transformations due to deforestation and climate change. Despite being species‐poor compared with central areas in Amazon or Cerrado, this ecotone harbors species with important adaptations that could hold the key to persistence in human‐disturbed landscapes or during periods of climate change.     

Leaf gas exchange,chlorophyll fluorescence and growth responses of <Emphasis Type="Italic">Melaleuca alternifolia</Emphasis> seedlings to flooding and subsequent recovery

Periodic flooding of trees in tropical floodplains and reservoirs where water levels fluctuate is a common phenomenon. The effects of flooding and subsequent recovery on gas exchange, chlorophyll fluorescence and growth responses of Melaleuca alternifolia seedlings, a tall shrub species used in floodplain and reservoir forest restoration in southern China, were studied during a grow season (from March to December in 2007). M. alternifolia seedlings were flooded for 180 days, drained and left to recover for another 60 days. Survival rates of the seedlings were 100% during the 180-day flooding period. Chlorophyll (Chl) content, net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) of the flooded seedlings were all significantly lower than those of the control. Significant reductions of photochemical quenching coefficient (q_p) and increases of nonphotochemical quenching (NPQ) in the flooded seedlings were observed. However, there were no significant differences in the maximal quantum yield of PSII photochemistry (F_v/F_m) between treatments. All seedlings survived during the two-month recovery period after the flooded treatment was drained, and the biomass and height of the recovered seedlings approached those of the control at the end of the experiment. During the first-month recovery period, Chl content, P _N, g _s and E in the recovered seedlings were all obviously low, then increased gradually and rose to the levels similar to the control by the end of the experiment. Quenching analysis revealed significant reductions of q_p and increments of NPQ in the recovered seedlings at the beginning of draining, and a nearly complete recovery for both parameters by the end of the experiment. However, F_v/F_m of the recovered seedlings did not differ significantly from the control during the recovery period. Our study demonstrated that M. alternifolia seedlings can survive and grow through 180 days of flooding with a subsequent 60-day recovery period in drained conditions, indicating that seedlings of this species would be suitable for afforestation in areas exposed to intermittent flooding.     

Response of wild mammals to seasonal shrinking-and-expansion of habitats due to flooding regime of the Pantanal, Brazil.

The Pantanal is a large savanna wetland (138,183 km2 in Brazil), important for its wildlife, fed by tributaries of the upper Paraguay River, center of South America (Brazil, touching Bolivia and Paraguay). Uplands are plateaus (250-1,200 m high, 215,000 km2 in Brazil) and flatland is the Pantanal (80-150 m high, 147,574 km2 in Brazil). Rivers are slow moving when they meet the flatland (slope 0.3-0.5 m/km east-west; 0.03-0.15 m/km north-south), periodically overflowing their banks, creating a complex seasonal habitat range. Recurrent shallow flooding occupies 80% of the Pantanal; during the dry season flooded areas dry up. Fluctuating water levels, nutrients and wildlife form a dynamic ecosystem. A flooding regime forms distinct sub-regions within the Pantanal. A mammal survey was carried out in the sub-region of the Rio Negro from April, 2003 through March, 2004 to study the diversity and abundance of terrestrial mammals during the dry and flooding seasons. A total of 36 species were observed in the field. The capybara Hydrochaeris hydrochaeris was the most frequent species, followed by the crab-eating-fox Cerdocyon thous and the marsh deer Blastocerus dichotomus. The highest abundance of species was observed during the dry season (August and September), when there is a considerable expansion of terrestrial habitats, mainly seasonally flooded grassland. Animal abundance (in terms of observed individual frequencies) varied during the dry and wet seasons and the seasonally flooded grassland was the most utilized habitat by mammals in the dry season.     

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.     

Structural and functional responses of plant communities to climate change‐mediated alterations in the hydrology of riparian areas in temperate Europe

The hydrology of riparian areas changes rapidly these years because of climate change‐mediated alterations in precipitation patterns. In this study, we used a large‐scale in situ experimental approach to explore effects of drought and flooding on plant taxonomic diversity and functional trait composition in riparian areas in temperate Europe. We found significant effects of flooding and drought in all study areas, the effects being most pronounced under flooded conditions. In near‐stream areas, taxonomic diversity initially declined in response to both drought and flooding (although not significantly so in all years) and remained stable under drought conditions, whereas the decline continued under flooded conditions. For most traits, we found clear indications that the functional diversity also declined under flooded conditions, particularly in near‐stream areas, indicating that fewer strategies succeeded under flooded conditions. Consistent changes in community mean trait values were also identified, but fewer than expected. This can have several, not mutually exclusive, explanations. First, different adaptive strategies may coexist in a community. Second, intraspecific variability was not considered for any of the traits. For example, many species can elongate shoots and petioles that enable them to survive shallow, prolonged flooding but such abilities will not be captured when applying mean trait values. Third, we only followed the communities for 3 years. Flooding excludes species intolerant of the altered hydrology, whereas the establishment of new species relies on time‐dependent processes, for instance the dispersal and establishment of species within the areas. We expect that altered precipitation patterns will have profound consequences for riparian vegetation in temperate Europe. Riparian areas will experience loss of taxonomic and functional diversity and, over time, possibly also alterations in community trait responses that may have cascading effects on ecosystem functioning.     

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.     

Seasonal flooding,topography, and organic debris interact to influence the emergence and distribution of seedlings in a tropical grassland

In seasonally flooded wetlands, inundation and associated organic debris deposition followed by a drawdown period can promote plant community diversity across space and time. Post‐flood regeneration might be influenced by the direct effect of flooding on seed dispersal and seedling emergence, as well as the indirect effect of organic debris on seed trapping and germination. Our objective was to examine the influence of seasonal flooding, topography, and organic debris cover on seedling distribution in a seasonally flooded grassland. We measured species richness, seedling abundance, and organic debris cover for 3 yr in a seasonally flooded grassland in the Pantanal, Brazil, at three topographic levels at the end of the flood season and during the dry season when there was no debris deposition. A total of 43 species were recorded, with no difference in species richness detected between seasons. However, the abundance of some species was higher post‐flood than during the dry period. The greatest seedling abundance and richness were found post‐flood at intermediate elevations, followed by high and the lowest elevations. Seed germination and seedling establishment were likely suppressed at low topographic positions due to shading from organic debris and poor drainage. Therefore, areas with predictable annual floods promote diversity by creating spatial and temporal variations in environmental conditions.     

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 anatomical adaptations induced by flooding in Cotula coronopifolia

Cotula coronopifolia is a wild annual Asteraceae that grows in periodically-flooded prone environments and seems highly tolerant to periodic flooding. Seedlings of about 15 cm were collected directly from the edge of Soliman sabkha (N-E Tunisia, semi-arid stage) and grown under greenhouse conditions. Two treatments were considered: drainage and flooding. After 56 days of treatment, flooded plants showed a pronounced growth increase. This performance was essentially associated with significant increment in biomass production of both shoots and roots (about 220% of the control). The appropriate response to flooding was also characterized by the ability of the species to maintain its water status under such conditions. Neither water content nor water potential showed a significant variation as compared to those of non-flooded plants. However, transpiration rate decreased slightly but significantly in flooded plants (from 0.86 to 0.64 mmol H2O m-2 s-1). Na+ and K+ concentrations were practically maintained under waterlogging conditions, except a significant increase of Na+ content in roots of flooded plants (157% of the control). These responses were concomitant with maintenance of photosynthetic rate. However, the contents of chlorophylls a and b increased to 167% and 295%, respectively. It seems that the enhancement in these photosynthetic pigments together with a significant improvement in water use efficiency (from 4.66 to 6.07 mmol CO2 mol-1 H2O) allowed to the species to compensate the decrease in photosynthetic rate. At the anatomical level, this species responded to flooding by a significant development of its root aerenchyma (+63%) and an increase in the lignification of its stem xylem tissues (+37%). Based on the presented data, the plant fitness under flooding conditions was a result of dynamic readjustment of several morphological, physiological, and anatomical adaptive traits. Flood requirement together with salt tolerance are responsible for the predominance of C. coronopifolia in a large area in its natural biotope where most plants cannot tolerate interactive effects of flooding and salinity.     

Seed bank of seasonally flooded grassland: experimental simulation of flood and post-flood

Wetland seed banks comprise the propagules of plant species that have species-specific germination requirements for germination in either flooded or dry conditions. At the community level, wetland structure and succession during and after a seasonal flooding event depends upon the early life-history requirements of species, including germination under flooded and dry conditions. We examined the effects of simulated flood and post-flood scenarios on seedling emergence from a seed bank of seasonally flooded grassland in the Pantanal, Brazil. Field samplings were conducted in both wet and dry seasons, both of which were subject to flood and post-flood conditions. A total of 70 species emerged from the seed bank, dominated by Poaceae and Cyperaceae. Sixteen species were exclusive to the wet and one exclusive to the dry season. The richness of perennial species was higher under flood conditions, while the richness of annuals was greater post-flood. In general, the aquatic and amphibious species exhibited a significant germination response to flooding. Terrestrial species only germinated in post-flood conditions, with higher richness in the dry season. Four species had high seedling abundance in both treatments. The capacity of regeneration by seeds is high in these grasslands and can be increased by seasonal flooding and drawdown. In these seasonally flooded grasslands, we observed three main germination strategies: under flooded conditions, aquatic and amphibious species; post-flood conditions, an explosion of annual amphibious and terrestrial species; and in moist soil, perennial terrestrial species. The differential responses to flooding versus post-flood conditions help to maintain the structure and species richness in the community over time.     

Does flooding of rice fields after cultivation contribute to wetland plant conservation in southern Brazil?

Question: Does flooding of rice fields after cultivation contribute to wetland plant conservation in southern Brazil? Location: Rice fields in the coastal plain of southern Brazil. Methods: Six rice fields with different management practices were randomly selected (three dry rice fields and three flooded rice fields). Six collections were carried out over the rice cultivation cycle. Richness and biomass were measured using the quadrat method. Results: A total of 88 macrophyte species was recorded. There was no statistical interaction between management practices and rice cultivation phases for macrophyte richness and biomass. Macrophyte species richness and biomass changed over time, but were similar between flooded and dry rice fields. The first three axes generated by detrended correspondence analysis explained 29% of the variation in species composition and the multivariate analysis of variance showed that there was a statistical interaction between management practices and agricultural periods. Conclusions: Rice fields may help to conserve an important fraction of the aquatic macrophyte diversity of wetlands of southern Brazil by providing the setting up of a greater number of species within the agricultural landscape. However, rice fields must not be viewed as surrogate systems for natural wetlands. The difference in species composition between flooded and dry rice fields is interesting in terms of biodiversity conservation. If rice producers could keep part of their agricultural land flooded during the fallow phase, this management practice could be an important strategy for the conservation of biodiversity in areas where natural wetlands have been converted to rice fields.     

Managed flooding can augment the benefits of natural flooding for native wetland vegetation

Managed flooding is increasingly being used to maintain and restore the ecological values of floodplain wetlands. However, evidence for its effectiveness is sometimes inconsistent and water available for environmental purposes often limited. We experimentally inundated a floodplain wetland (or “billabong”) in late spring by pumping water from its adjacent creek, aiming to promote the native wetland flora and suppress terrestrial exotics. Vegetation was surveyed before (spring) and after (late summer) the managed flood in the experimental billabong and in three control billabongs. Floodplain water levels were continuously monitored. Wet conditions caused two of the control billabongs to also flood, but to a lesser extent than the experimental billabong. We therefore assessed vegetation changes relative to flooding duration. With increasing flooding duration, the cover of wetland vegetation (amphibious and aquatic species) increased and the cover of terrestrial and exotic vegetation decreased, with these effects largest in the deliberately flooded billabong. Flooding durations greater than 20 days generally resulted in increased cover of wetland plants and restricted the growth of terrestrial plants. Reinstatement of more appropriate flooding regimes can thus promote native wetland plants, while suppressing terrestrial exotic species. Our study also provides evidence for the use of modest water allocations to augment the benefits of natural flooding in the maintenance and restoration of native wetland plant communities.     

Maintaining and restoring hydrologic habitat connectivity in mediterranean streams: an integrated modeling framework

Hydrologic alterations designed to provide a stable water supply and to prevent flooding are commonly used in mediterranean-climate river (med-rivers) basins, and these alterations have led to habitat loss and significant declines in aquatic biodiversity. Often the health of freshwater ecosystems depends on maintaining and recovering hydrologic habitat connectivity, which includes structural components related to the physical landscape, functionality of flow dynamics, and an understanding of species habitat requirements for movement, reproduction, and survival. To advance our understanding of hydrologic habitat connectivity and benefits of habitat restoration alternatives we provide: (1) a review of recent perspectives on hydrologic connectivity, including quantitative methods; and (2) a modeling framework to quantify the effects of restoration on hydrologic habitat connectivity. We then illustrate this approach through a case study on lateral hydrologic habitat connectivity that results from channel restoration scenarios using scenarios with different historic and climate-change flows to restore fish floodplain habitat in a med-river, the San Joaquin River, California. Case study results show that in addition to the channel alterations, higher flows are required to recover significant flooded habitat area, especially given reductions in flows expected under climate change. These types of studies will help the planning for restoration of hydrologic habitat connectivity in med-rivers, a critical step for mediterranean species recovery.     

Effect of soil flooding on photosynthesis,carbohydrate partitioning and nutrient uptake in the invasive exotic Lepidium latifolium

Lepidium latifolium L. is an invasive exotic crucifer that has spread explosively in wetlands and riparian areas of the western United States. To understand the ecophysiological characteristics of L. latifolium that affect its ability to invade riparian areas and wetlands, we examined photosynthesis, chlorophyll concentration, carbohydrate partitioning and nutrient uptake in L. latifolium in response to soil flooding. Photosynthesis of flooded plants was about 60–70% of the rate of unflooded controls. Chlorophyll concentrations of flooded plants were about 60–70% of the unflooded plants during 15–50 days of flooding. Flooding resulted in an increase in leaf starch concentration, but root starch concentration was not significantly affected. However, concentrations of soluble sugar were significantly higher in both leaves and roots of flooded plants than unflooded controls. On day 50 after initial flooding, the concentrations of N, P, K and Zn in leaves of flooded plants were lower than in control plants. The concentrations of Mn and Fe in leaves of flooded plants were eight and two times those of control plants, respectively. In contrast, N, P, K and Zn concentrations of roots of flooded plants were slightly higher than in unflooded plants. The concentrations of Fe and Mn in roots of flooded plants were 15 and 150 times those of the control plants, respectively. The transport of P, K, and Zn to shoots decreased and that of Mn increased under flooding. The accumulation of N, K and Zn in roots decreased and that of Mn increased in response to flooding. The results suggested that the maintenance of relatively high photosynthesis and the accumulation of soluble sugar in roots of flooded plants are important adaptations for this species in flooded environments. Despite a reduction in photosynthesis and disruption in nutrient and photosynthate allocation in response to flooding, L. latifolium was able to survive 50 days of flooding stress. Overall, L. latifolium performed like a facultative hydrophyte species under flooding.     

Role of Ethylene in Induction of Flooding Damage in Sunflower

The possibility that symptoms of flooding damage in plants are primarily caused by an accumulation of ethylene was investigated using pot-grown sunflower (Helianthus annuus) plants. When plants were flooded to the basal pairs of leaves, ethylene in roots and stems below the water line began to increase. This coincided with the start of hypocotyl hypertrophy and new root formation in hypocotyls, which continued for 14-16 days. There were highly significant correlations between ethylene concentration and number of roots and hypocotyl diameter. After approximately 4 days of flooding, ethylene concentrations in stems between nodes for the 1st and 3rd basal pairs of leaves started to increase, coinciding with initiation of chlorophyll breakdown and epinasty of the 2nd basal pairs of leaves. Thus, there were correlations between ethylene concentration and chlorophyll breakdown and epinasty. The lower the leaves, the more chlorophyll breakdown among 1st, 2nd, 3rd, and 4th basal pairs of leaves. The longer the flooding, the more severe the flooding damage; and even when returned to normal condition, plants flooded longer than 3 days were not able to recover from flooding damage. A gas chromatographic study revealed that Ethephon was absorbed by roots and decomposed to ethylene in the plant. Damage symptoms caused by soil application of Ethephon, such as reduced stem height, chlorophyll breakdown, epinasty of the 2nd basal pairs of leaves, and hypocotyl hypertrophy, were almost identical with those caused by soil flooding treatment. Microscopic studies revealed that radially enlarged cells and increased intercellular spaces in the cortex were the major contribution to the increased hypocotyl diameter in both flooded and Ethephon-treated plants. It is concluded that the increase in ethylene concentration in flooded plants is largely, although not exclusively, responsible for flooding damage symptoms.     

Interactive effect of flooding and grazing on the growth of Serengeti grasses

Summary Grazing and flooding may potentially interact in particular habitats of many grassland regions around the world. We tested the hypothesis that grazing and flooding induce different and largely opposed allocation responses in individual plants. As a result, their combined effect on plant growth would be negative. We studied the response of three grass species from the Serengeti ecosystem (Tanzania) to the effects of flooding and clipping. Plants under the combined effect of flooding and clipping had lower growth rates than plants growing under the effect of either of the two factors individually. Plants under flooding grew taller and allocated more resources to stem growth than controls; for two of the three species, flooded plants also generated a new root system above soil level. All these morphological and physiological responses conflict with the ability of a plant to respond to defoliation with minimum reduction in growth rates. The three species showed a response to flooding reflecting their distribution ranges in the field: the species from the most flood-prone habitat showed a positive effect of flooding on growth, whereas the species from dry uplands showed a strong negative effect of flooding. Flood-tolerant species were taller and less tolerant of clipping than flooding sensitive species. Our results suggest that, in ecological time, individuals subjected to both flooding and grazing have their growth reduced to a greater extent than by either of the two factors acting individually, whereas in evolutionary time, species adapted to flooding are poor grazing tolerators and species adapted to grazing are poor flooding tolerators.