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

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

The Effects of Intermittent Flooding on Seedlings of Three Forest Species

Under greenhouse conditions, seedlings of three forest species, baldcypress (Taxodium distichum), nuttall oak (Quercus nuttallii), and swamp chestnut oak (Quercus michauxii) were subjected to an intermittent flooding and subsequent physiological and growth responses to such conditions were evaluated. Baldcypress showed no significant reductions in stomatal conductance (g _s) or net photosynthetic rate (P _N) in response to flood pulses. In nuttall oak seedlings g _s and P _N were significantly decreased during periods of inundation, but recovered rapidly following drainage. In contrast, in swamp chestnut oak g _s was reduced by 71.8 % while P _N was reduced by 57.2 % compared to controls. Baldcypress displayed no significant changes in total mass while oak species had significantly lower leaf and total mass compared to their respective controls. Thus baldcypress and nuttall oak showed superior performance under frequent intermittent flooding regimes due to several factors including the ability for rapid recovery of gas exchange soon after soil was drained. In contrast, swamp chestnut oak seedlings failed to resume gas exchange functions after the removal of flooding.     

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

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

Effects of soil flooding and changes in light intensity on photosynthesis of <Emphasis Type="Italic">Eugenia uniflora</Emphasis> L. seedlings

The increased frequency of heavy rains as a result of global climate change can lead to flooding and changes in light availability caused by the presence of thick clouds. To test the hypothesis that reduction in light availability can alleviate the harmful effects of soil flooding on photosynthesis, the authors studied the effects of soil flooding and acclimation from high to low light on the photosynthetic performance of Eugenia uniflora. Seedlings acclimated to full sunlight (about 35 mol m⁻² d⁻¹) for 5 months were transferred to partial sunlight (about 10 mol m⁻² d⁻¹) and were either subjected to soil flooding or not flooded. Chlorophyll fluorescence was measured throughout the flooding period and leaf gas exchange was measured 16 days after flooding was initiated. Minimal fluorescence yield (Fo) was significantly higher and the quantum efficiency of open PSII centres (Fv/Fm) was significantly lower in flooded than in non-flooded plants in full sunlight. Sixteen days after flooding was initiated, stomatal conductance (gs_sat) and net photosyntheses expressed on a leaf area (A_sat-area), weight (A_sat-wt) and chlorophyll (A_sat-Chl) basis decreased in response to soil flooding. Flooding decreased stomatal conductance by similar amounts in full and partial sunlight, but A_sat-area in partial and full sunlight was 3.4 and 16.8 times lower, respectively, in flooded than in non-flooded plants. These results indicate that changes from full to partial sunlight during soil flooding can alleviate the effects of flooding stress on photosynthesis in E. uniflora seedlings acclimated to full sunlight. The responses of photosynthesis in trees to flooding stress may be dependent on changes in light environment during heavy rains.     

Responses of Melaleuca quinquenervia seedlings to flooding

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

Stomatal Responses of Fraxinus pennsylvanica Seedlings during and after Flooding

Effects of flooding for 10, 20, 30, or 40 days on leaf diffusion resistance (r₁) of fraxinus pennsylvanica seedlings were studied during the period of flooding and continuing for 17 days after flooding. All Flooding treatments induced stomatal closure, as indicated by increased r₁. There was some evidence of stomatal adaptation to flooding, with stomata beginning to reopen after a critical period of flooding. After termination of flooding, stomata opened further within 6 to 10 days to near preflooding levels. The degree of stomatal opening was only slightly higher after 10 days than after 40 days of flooding. Some stomata may have been permanently damaged by flooding. The stomatal adaptation to flooding as well as rapid recovery of stomatal opening, even after prolonged flooding, appeared to be important factors in the flooding tolerance of Fraxinus petmsylvanica and are consistent with its distribution on wet sites.     

Changes in photosynthesis and fluorescence in response to flooding in emerged and submerged leaves of Pouteria orinocoensis

In the seasonally flooded forest of the Mapire River, a tributary of the Orinoco, seedlings remain totally covered by flood water for over six months. In order to characterize the physiological response to flooding and submergence, seedlings of the tree Pouteria orinocoensis, an important component of the forest vegetation, were subjected experimentally to flooding. Flooding was imposed gradually, the maximum level of flood including submerged and emerged leaves. After 45 d a severe reduction of net photosynthetic rate (P _N) and stomatal conductance (g _s) was observed in emerged leaves, whereas leaf water potential remained constant. The decrease in P _N of emerged leaves was associated to an increase in both relative stomatal and non-stomatal limitations, and the maintenance of the internal/air CO₂ concentration (C _i/C _a) for at least 20 d of flooding. After this time, both P _N and g _s became almost zero. The decrease in photosynthetic capacity of emerged leaves with flooding was also evidenced by a decrease in carboxylation efficiency; photon-saturated photosynthetic rate, and apparent quantum yield of CO₂ fixation. Oxygen evolution rate of submerged leaves measured after three days of treatment was 7 % of the photosynthetic rate of emerged leaves. Submersion determined a chronic photoinhibition of leaves, viewed as a reduction in maximum quantum yield in dark-adapted leaves, whereas the chlorophyll fluorescence analysis of emerged leaves pointed out at the occurrence of dynamic, rather than chronic, photoinhibition. This was evidenced by the absence of photochemical damage, i.e. the maintenance of maximum quantum yield in dark-adapted leaves. Nevertheless, the observed lack of complementarity between photochemical and non-photochemical quenching after 12 d of flooding implies that the capacity for photochemical quenching decreased in a non-co-ordinate manner with the increase in non-photochemical quenching.     

Soil temperature and flooding effects on two species of citrus

Summary Rough lemon (Citrus jambhiri Lush.) and sour orange (C. aurantium L.) seedlings were grown at constant soil temperatures of 16, 24, and 33 C for 3 months. Shoot and root growth of rough lemon was greatest at 33 C while growth of sour orange was greatest at 24 C. There were no significant effects of soil temperature on shoot: root ratio, leaf water potential or stomatal conductance. The hydraulic conductivity of intact root systems of both species was highest when seedlings were grown at 16 C. Thus, acclimation through greater root conductivity at low soil temperature may have compensated for decreased root growth at 16 C and negated effects of soil temperature on plant water relations. Half the plants growing at each soil temperature were subsequently flooded. Within 1 week, the soil redox potential (Eh) dropped below zero mV, reaching a minimum Eh of –250mV after 3 weeks of flooded conditions. Flooded plants exhibited lower root conductivity, a cessation of shoot growth, lower leaf water potentials, lower stomatal conductances, and visual sloughing of fibrous roots. Decreases in root conductivity in response to flooding were large enough to account for the observed decreases in stomatal conductance.Florida Agricultural Experiment Stations Journal Series No. 4080.     

Salinity and flooding stress effects on mycorrhizal and non-mycorrhizal citrus rootstock seedlings

Seedlings of the rootstocks Pineapple sweet orange (SwO), Carrizo citrange (CC), and sour orange (SO) were grown in low phosphorus (P) sandy soil and either inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus,Glomus intraradices, or were non-mycorrhizal (NM) and fertilized with P. VAM and NM seedings of similar shoot size and adequate P-status were selected for study of salinity and flooding stress. One-third of each of the VAM and NM plants were given 150 mM NaCl for a period of 24 days. One-third of the plants were placed into plastic bags and flooded for 21 days while the remaining third were non-stressed controls. In general, neither stress treatment affected mycorrhizal colonization. Salinity stress reduced the hydraulic conductivity of roots, leaf water potential, stomatal conductance and net assimilation of CO₂ (ACO₂) of mycorrhizal and non-mycorrhizal seedlings to a similar extent. VAM plants of CC and SO accumulated more Cl in leaves than NM plants. Cl was higher in non-mycorrhizal roots of SwO and CC than in mycorrhizal roots. Flooding the root zone for 3 weeks did not produce visible symptoms in the shoot but did influence plant water relations and reduce ACO₂ of all 3 rootstocks. VAM and NM plants of each rootstock were affected similarly by flooding. Comparable reduction in nitrogen and P content of both mycorrhizal and non-mycorrhizal plants suggested that flooding stress was primarily affecting root rather than hyphal nutrient uptake. Florida Agricultural Experimental Station Journal Series No. 7773.     

Variations among Woody Angiosperms in Response to Flooding

Effects of flooding on young Populus deltoides, Salix nigra, Eucalyptus camaldulensis, E. globulus, Ulmus americana, Quercus rubra and Fraxinus pennsylvanica plants were studied. Flooding variously induced several sequential physiological disturbances, with stomatal closure among the earliest responses. Subsequent responses included inhibition of root growth, alterations in root and stem morphology, formation of adventitious roots, and leaf senescence. In amphistomatous species (Populus deltoides, Salix nigra, Eucalyptus camaldulensis) flooding rapidly induced stomatal closure on the adaxial leaf surface. It also significantly induced stomatal closure on the abaxial epidermis of Populus deltoides but not that of Salix nigra or Eucalyptus camaldulensis. In hypostomatous species (Eucalyptus globulus, Ulmus americana) flooding significantly induced stomatal closure on the abaxial surface within 3 days. Stomatal responses to flooding were not correlated with leaf water stress. In both long- and short-term experiments, flooding did not significantly increase plant water stress. These results deemphasized the importance of plant water stress in inducing plant responses to flooding. The importance of various hormones in inducing flooding symptoms is discussed.     

Developing a mathematical model for variation of physiological responses of Jatropha curcas leaves depending on leaf positions under soil flooding

The effect of soil flooding on photosynthesis, transpiration and stomatal conductance of Jatropha curcas seedlings were studied under natural environmental variables. Soil flooding reduced photosynthesis (P _N), transpiration (E) and stomatal conductance (gs) in response to leaf positions of Jatropha curcas plants. Based on the results, we conclude that decrease in stomatal opening and stomatal limitation of photosynthesis, followed by decrease in individual leaf area are the main causes of reductions in carbon uptake of flooded seedlings. A mathematical relationship was successfully developed to describe photosynthesis, transpiration and stomatal response of Jatropha under soil flooding stress.     

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.     

The effect of Ca2+-stress on fluxes of Ca2+ and K+ in wheat and cucumber

Betula papyrifera Marsh, seedlings adapted very poorly to flooding for up to 60 days. Responses to flooding included increased ethylene production; stomatal closure; leaf senescence; drastic inhibition of shoot growth, cambial growth, and root growth; decay of roots, and death of many seedlings. Flooding inhibited growth of leaves that formed prior to flooding, inhibited formation of new leaves, and induced abscission of old leaves. As a result of extensive leaf abscission, fewer leaves were present after flooding than before flooding was initiated. The drastic reduction in leaf area was associated with greatly decreased growth of the lower stem and roots. No evidence was found of adaptive morphological changes to flooding. The data indicate that intolerance of B. papyrifera seedlings to flooding is an important barrier to regeneration of the species on sites subject to periodic inundation.     

Gas exchange and photosynthesis of Eucalyptus camaldulensis seedlings inoculated with different ectomycorrhizal symbionts

Eucalyptus camaldulensis Dehnh. seedlings inoculated with Pisolithus tinctorius (Pers.) Coker & Couch and Thelephora terrestris Ehrl. per Fr. were grown in well watered soil (_s –0.03 MPa) or subjected to a long-term soil water stress of up to –1.0 MPa over 13-week period in a glasshouse. After 13 weeks, all seedling containers were watered to field capacity and then water was withheld from the E. camaldulensis seedlings to induce a short-term drought. Diurnal measurements of seedling photosynthesis rate (A), leaf stomatal conductance (g) and leaf water potential (_p) were completed before, during, and after the short term drought. Although they were growing in an equal soil volume, photosynthesis rate (A), leaf stomatal conductance and leaf water potential (_p) of larger seedlings with P. tinctorius ectomycorrhizae were similar to those of smaller seedlings colonized with T. terrestris during the short-term drought period. Seedlings inoculated with Pisolithus tinctorius maintained higher photosynthesis rates over the course of the short-term drought. Thus, P. tinctorius ectomycorrhizae appear to be more efficient than those of T. terrestris in assisting seedlings to maintain gas exchange and photosynthesis under limited soil moisture conditions.     

Effects of flood pre-conditioning on responses of three bottomland tree species to soil waterlogging

The potential for improving the performance of seedlings of woody species under flood stress was evaluated. Seedlings of baldcypress (Taxodium distichum), nuttall oak (Quercus nuttallii), and swamp chestnut oak (Quercus michauxii) were subjected to a two-phase study in which soil moisture regime was controlled. During Phase I, plants were randomly assigned to either: Control, well-watered and drained conditions; or intermittent flooding, alternating flooding and drained conditions. Following completion of Phase I, seedlings in each treatment were assigned randomly to one of two treatments: well-watered but drained or continuously flooded. Thus during Phase II, plants were divided into four groups: Control (A), intermittent flooding during Phase I and draining during Phase II (B), control during Phase I and continuously flooding during Phase II (C), and intermittently flooded during Phase I and continuously flooded during Phase II (D). Stomatal conductance (gw) in baldcypress increased in flooded treatments ranging from 112 to 128 percnt; of controls. Net photosynthesis (Pn) in baldcypress was not significantly affected by flooding while in oaks, Pn and gw decreased significantly in response to flooding. In oaks, flood pre-conditioning did not affect growth or physiological responses during phase II significantly. However, stomatal conductance of flood-hardened baldcypress was increased compared to seedlings that were not hardened. The study species developed significantly higher root porosity under flooded conditions compared to controls. Flood pre-conditioning did not appear to increase root porosity although it may have accelerated the process.     

The impact of flooding and drought on seeds of Cnidium dubium, Gratiola officinalis, and Juncus atratus, three endangered perennial river corridor plants of Central European lowlands

The effects of flooding and drought on the post-flooding survival and germinability of seeds in the soil were investigated for three endangered river corridor angiosperms Cnidium dubium (Schkuhr) Thell., Gratiola officinalis L. and Juncus atratus Krocker. Freshly matured seeds were placed in small nylon bags and were buried in pots filled with soil collected from a wetland along the Lower Havel River. The pots were subjected to different winter/spring flooding treatments (30, 60, 120, and 180 days) and to drought for up to 3 years. Every year in spring, after drawdown, the seeds were exhumed, counted and tested for germination. Seed mortality of flooded seeds was low, except for C. dubium. Flooding did not affect germination of seeds of J. atratus. Flooding of seeds of C. dubium and G. officinalis had inconsistent, but no beneficial effects on germination of either. The year of harvest and the interannual variability of weather are found to be the main factors affecting germinability of C. dubium and G. officinalis in the course of the three consecutive years.The experiment showed that the capacity of the flooded seed bank to maintain germination ability and viability for extended periods of time could be directly related to the flooding duration of natural habitats of the three species. The three species are resilient to variable flooding. Seeds appeared desiccation tolerant, and single flooding events do not deplete their seed bank.     

Seedling growth of Mora gonggrijpii,a large seeded climax species,under different soil and light conditions

Mora gonggrijpii seedlings were harvested on two soil types in small gaps and in the forest understorey. The seedlings appear to be intolerant of lateritic soil with poor drainage either as a result of poor aeration or reduction of the soil which is rich in iron and aluminium. Biomass allocation is not greatly affected on the lateritic soil as compared to the better drained sandy soil. During the initial two months light has no effect on the growth of the seedlings but soil type does. In the same period cotyledon depletion is equal, suggesting higher turnover rates or respiration rates on the poorly drained lateritic soil. From the second to the seventh month light stimulates growth, but Mora gonggrijpii seedlings are not plastic in their morphology. Leaf area remains constant during this period, and no branching is observed in the higher light environment. Specific leaf area, leaf area ration, net assimilation rate and relative growth rate are all among the lowest thus far reported for tree seedlings. Evidence is presented that these characteristics are found often in large seeded (climax) species.     

Which seed origin provides better tolerance to flooding and drought when restoring to face climate change?

Our goal was to establish the tolerance to flooding and drought of seedlings from a hydric gradient of different seed sources to provide recommendations for forest restoration in the face of climate change. We used Drimys winteri var. chilensis, a tree species that grows from extreme arid zones to wetlands along Chile, as the study subject. We expected that seedlings of xeric origin would perform better in drought conditions than populations from moist environments, and vice versa for flooding tolerance. We collected D. winteri seeds from xeric, mesic and wet environments. Seedlings at two development stages were submitted to an extreme flooding and drought treatment during 2 or 4 months in a common garden. After the flooding and drought assays finished, the number of surviving and damaged seedlings, lenticels and adventitious root presence, height, new leaves and specific leaf area, shoot/root ratio, water potential and/or chlorophyll fluorescence (Fv/Fm), were recorded. We found that flooding and drought affected almost all the parameters studied negatively. The xeric population seedlings, at both development stages studied, were the most tolerant to the drought and, unexpectedly, also to the flooding treatment. We recommend restoring with seedlings of xeric origin especially in arid areas where sudden flooding is frequent, as occurs in the Andes Mountains. In the face of climate change, we recommend carrying out common garden and field studies before advising which population origin should be used for restoration, since they do not always respond in accordance with expected patterns of local adaptation.     

Chemical and Hydraulic Influences on the Stomata of Flooded Plants

Flooding the soil greatly reduced stomatal opening and leafgrowth rate of pea (Pisum sativum L cv. Feltham First) seedlings.This was despite the fact that leaf water potential and turgorwere not significantly affected by the treatment. Potassiumuptake and transport to the leaves was reduced by flooding.Stomata of flooded plants could be reopened by incubating leavesin solutions containing KC1. These observations raise the possibilitythat nutrient deficiency may limit stomatal opening and growthin flooded plants. We also consider the possibility that potassiumdeficiency may interact with a modification in the balance ofgrowth regulators in the leaves to modify stomatal behaviourand growth. Key words: Flooding, stomata, leaf growth, water relations, potassium     

Structure and regeneration patterns of the <Emphasis Type="Italic">Juniperus excelsa</Emphasis> Bieb. stands in the central part of the Nestos valley in the northeast of Greece,in the context of anthropogenic disturbances and nurse plant facilitation

In order to analyse the structure and regeneration patterns of the Juniperus excelsa Bieb. stands on the slopes of the Nestos valley, 40 sample plots were established and more than 120 J. excelsa trees and seedlings were cut and collected. The main results of this research indicate that stand structure, species mixture, age structure and stand development patterns were strongly determined by the interaction of anthropogenic disturbances such as grazing and illegal cuttings, and the facilitation of adult J. excelsa trees in the establishment of the J. excelsa seedlings. Their effect was influenced by site quality and J. excelsa together with other species ecology. Under present conditions, almost all J. excelsa seedlings, except for a small number of plants, are found under the facilitation of nurse plants that belong to the same species. It seems that the layer of litter underneath the nurse plants combined with the protection of seedlings against grazing play an important role in the establishment and growth of J. excelsa seedlings.