Effects of oil on internal gas transport, radial oxygen loss, gas films and bud growth in Phragmites australis

Background and Aims

Oil pollution of wetlands is a world-wide problem but, to date, research has concentrated on its influences on salt marsh rather than freshwater plant communities. The effects of water-borne light oils (liquid paraffin and diesel) were investigated on the fresh/brackish wetland species Phragmites australis in terms of routes of oil infiltration, internal gas transport, radial O₂ loss (ROL), underwater gas films and bud growth.

Methods

Pressure flow resistances of pith cavities of nodes and aerenchyma of leaf sheaths, with or without previous exposure to oil, were recorded from flow rates under applied pressure. Convective flows were measured from living excised culms with oiled and non-oiled nodes and leaf sheaths. The effect of oil around culm basal nodes on ROL from rhizome and root apices was measured polarographically. Surface gas films on submerged shoots with and without oil treatment were recorded photographically. Growth and emergence of buds through water with and without an oil film were measured.

Key Results

Internodes are virtually impermeable, but nodes of senesced and living culms are permeable to oils which can block pith cavity diaphragms, preventing flows at applied pressures of 1 kPa, natural convective transport to the rhizome, and greatly decreasing ROL to phyllospheres and rhizospheres. Oil infiltrating or covering living leaf sheaths prevents humidity-induced convection. Oil displaces surface gas films from laminae and leaf sheaths. Buds emerge only a few centimetres through oil and die.

Conclusions

Oil infiltrates the gas space system via nodal and leaf sheath stomata, reducing O₂ diffusion and convective flows into the rhizome system and decreasing oxygenation of phyllospheres and rhizospheres; underwater gas exchange via gas films will be impeded. Plants can be weakened by oil-induced failure of emerging buds. Plants will be most at risk during the growing season.Key words: Phragmites australis, oil pollution, convective flow, pressure flow resistance, phyllosphere oxygenation, rhizosphere oxygenation, underwater gas films, bud emergence, stomata, pith cavity diaphragms, leaf sheath aerenchyma, rhizome aeration     

Rice: sulfide-induced barriers to root radial oxygen loss, Fe2+ and water uptake, and lateral root emergence

BACKGROUND AND AIMS: Akagare and Akiochi are diseases of rice associated with sulfide toxicity. This study investigates the possibility that rice reacts to sulfide by producing impermeable barriers in roots. METHODS: Root systems of rice, Oryza sativa cv. Norin 36, were subjected to short-term exposure to 0.174 mm sulfide (5.6 ppm) in stagnant solution. Root growth was monitored; root permeability was investigated in terms of polarographic determinations of oxygen efflux from fine laterals and the apices of adventitious roots, water uptake, anatomy and permeability to Fe2+ using potassium ferricyanide. KEY RESULTS: Both types of root responded rapidly to the sulfide with immediate cessation of growth, decreased radial oxygen loss (ROL) to the rhizospheres and reduced water uptake. Profiles of ROL measured from apex to basal regions of adventitious roots indicated that more intense barriers to ROL than normal were formed around the apices. Absorption of Fe2+ appeared to be impeded in sulfide-treated roots. In adventitious roots, deposition of lipid material (suberisation) and thickenings of walls within the superficial cell layers were obvious within a week after lifting the treatment and could prevent the emergence of laterals and commonly result in their upward longitudinal growth within the cortex. Death of laterals sometimes occurred prior to emergence; emergent laterals eventually died. In adventitious roots, blockages formed within the vascular and aeration systems in response to the sulfide. CONCLUSIONS: In both adventitious and lateral roots, sulfide-induced cell wall suberization and thickening of the superficial layers were correlated with reduced permeability to O2, water and Fe2+. This study sheds light on some of the symptoms of diseases such as Akiochi. The results correlate with the authors' previous findings on the effects on roots of sulfide and lower organic acids in Phragmites and of acetic acid in rice.     

Effects of lead contamination on the clonal propagative ability of Phragmites australis (common reed) grown in wet and dry environments

Clonal propagation is important for the survival and maintenance of the common reed Phragmites australis. Pot culture experiments were conducted to investigate the effects of lead (Pb) concentration (0, 500, 1500, 3000, 4500 mg·kg^?1) and water stress on the clonal reproductive ability of this species. The Pb concentration found in plant organs, in decreasing order, was roots >shoots >rhizomes. There was a negative relationship between the growth of clonal propagative modules (excluding axillary shoot buds) and Pb concentrations, which caused a decrease in biomass, rhizome growth and number of axillary and apical rhizome buds. Daughter axillary shoots exhibited a tolerance strategy, with no significant change in their number; the axillary and apical rhizome buds, daughter apical rhizome shoots and rhizomes exhibited compensatory growth during the late stage of Pb (excluding 4500 mg·kg^?1) treatment in a wet environment. Pb applications above 500 mg·kg^?1 reduced these parameters significantly in the drought treatment, except for the number of axillary shoot buds, which did not change. Our results indicate that clonal propagative resistance to Pb contamination can occur via tolerance strategies, compensatory growth and a Pb allocation strategy, enabling these reeds to maintain population stability in wet environments. However, clonal modular growth and reproductive ability were inhibited significantly by the interaction between drought and Pb, which would cause a decline in P. australis populations in a dry environment. Lead concentrations of 4500 and 500 mg·kg^?1 in soils might meet or exceed the Pb tolerance threshold of clonally propagated reeds in wet and dry environments, respectively.     

CONTROL OF SHOOT-RHIZOME DIMORPHISM IN THE WOODY MONOCOTYLEDON,CORDYLINE (AGAVACEAE)

In Cordyline terminalis negatively geotropic leafy shoots and positively geotropic rhizomes develop from single axillary buds on either shoots or rhizomes. All axillary buds have similar morphogenetic potential when released from apical dominance. Experiments in which the orientation of the apex is changed, organs removed, or growth regulators applied indicate that after a rhizome is initiated, it is maintained as a rhizome by auxin originating in the leafy shoot. When auxin levels are lowered by changes in the orientation of the axis or shoot removal, the rhizome apex becomes a shoot apex, which appears to be the stable state of the actively growing apex. Benzyl adenine when applied exogenously to the apex or lateral buds has the same effect as lowering the auxin level. Gibberellic acid has no effect on the apex or lateral buds. High levels of exogenous naphthaleneacetic acid cause bud release and development of rhizomes from previously inhibited axillary buds of the shoot. However, it was not possible to convert a shoot apex into a rhizome apex by auxin treatment. It is suggested that the release of buds on the lower side of horizontal branches and of buds directly above a stem girdle is caused by high auxin levels on the lower side or distal to the girdle. The experimental results are discussed in relation to naturally occurring shoot-rhizome dimorphism.     

Rice and Phragmites: effects of organic acids on growth, root permeability, and radial oxygen loss to the rhizosphere

Young Phragmites plants were grown in two cocktails of monocarboxylic acids (C(1)-C(5)) at pH 6, where the concentration of each acid was innocuous and the total undissociated (potentially toxic) concentrations were 0.35 mmol/L and 0.42 mmol/L. Rice plants were subjected to 1.5 mmol/L acetic acid at pH 4.5 (undissociated concentration = 1.05 mmol/L). In Phragmites, each cocktail curtailed root growth especially and induced premature shoot senescence. In both species, after 3-5 d of treatment, radial oxygen loss (ROL) from apical regions of adventitious roots, and from Phragmites laterals, was reduced to very low values and associated with cell wall lignification and suberization in the surface cell layers. At later stages of treatment, rice responded to acetic acid in similar ways to Phragmites, with the development of intercellular and callus type occlusions in the gas space system, vascular blockages, and the failure of laterals to emerge. The results are relevant to the supply of oxygen from Phragmites roots to sediments for the phytopurification of waste waters, to the efflux of methane and carbon dioxide from wetlands, and to rice cultivation.     

The Structure,Elongation and Thickening of Rhizome in Nelumbo nucifera Gaertn

The seedling of Nelurnbo nucifera is erect and its internodes are very short with four Alternately arranged floating leaves. During the juvenile stage, the shoot elongates remarkably and forms the horizontal rhizome. Each leaf grows out from the dorsal side of the node of the rhizome. There are two kinds of terminal buds in the juvenile shoot. (1) vegetative bud and (2) mixed bud. The axillary scale is the derivative part of the leaf. It forms an ochrea around the terminal bud. The winter buds on the annual shoot are all mixed buds. The vessels are absent in the rhizome and no cambium exists. During tile early growth of the rhizome, the rib meristems contribute mainly to the internode elongation. Later however, divisions are seen to commence in the parenchymatous tissue of the internode. As a result of these divisions the internode becomes elongated. The tuberization of the rhizome is built up from cell divisions of three kinds of tissues: (1) primary thickening meristems, (2) cells of the vascular bundles and (3) parenchyma of cortex. But, the growth in thickness of the rhizome seems to be chiefly due to the enlargement of parenchymatous cells.     

Resistance to freezing in liquid nitrogen of carnation (Dianthus caryophyllus L. var Eolo) apical and axillary shoot tips excised from different aged in vitro plantlets

The ability of shoot tips from carnation (Dianthus caryophyllus L., var. Eolo) cultured in vitro to develop resistance to freezing in liquid nitrogen depends on the physiological state of the cell material and the pretreatment conditions. Regrowth rates close to 100% have been obtained with apical shoot tips isolated from 2 month-old stems, precultured on medium supplemented with sucrose (0.75M) and treated with dimethylsulfoxide (5% or more). Resistance of axillary shoot tips decreased progressively as a funtion of their distance from the apical shoot tip. During the development of the stem from axillary buds (obtained by cutting), progressive increases in the regrowth rate of frozen apices were noted, from 30% before cutting (axillary buds) to 98% after 3 weeks of culture.Abbreviations DMSO dimethylsulfoxide - LN liquid nitrogen     

Differences in rhizome aeration of Phragmites australis in a constructed wetland

Waterlogged soils are a challenging environment for plants due to anoxic conditions and enrichment of phytotoxic substances in the rhizosphere. High contents of organic matter in the substrate as present in constructed wetlands (CW) may amplify these effects. The influence of substrate organic matter on the rhizome aeration of Phragmites australis (common reed) was investigated in a CW for sewage treatment on two sites with different organic concentrations: (a) inflow part of the bed with a high organic charge (high-organic site) and (b) outflow zone of the bed with a lower organic charge (low-organic site). Several diurnal oxygen (O₂) courses were recorded inside the rhizomes using micro-optodes. Maximum O₂ percentages in the rhizomes were generally the same at both sites, but minimum O₂ values were lower in the low-organic site than in the high-organic site, leading to higher amplitudes at the low-organic site. The results suggest that oxygen release from the roots is hampered under high-organic compared to low-organic conditions. This observation might be explained by changes in gas conductivity of the roots, but also by alterations in microbial oxygen demand, under different organic burden.     

THE FATE OF THE DWARF SHOOT APEX IN BRISTLECONE PINE (PINUS LONGAEVA)

The fate of the pine dwarf shoot (DS) apex after needle initiation has been controversial. Dwarf shoot primordia of Pinus longaeva were examined to determine the developmental basis for DS with and without interfoliar buds. Interfoliar buds are microscopic buds derived from the original terminal apex of the DS. In October, all the DS primordia are similar in size and appearance. However, as the needles elongate in the following June the apices of more proximal DS decrease in size, such that by July there is a clear diminishing size gradient of apical domes in going from the most distal to the most proximal positions. The distal DSs start to form bud scales in July and have fully formed interfoliar buds by mid-August. In contrast, those DS apices lacking protective bud scales at needle maturity become suberized and can never proliferate into long shoots. The distal placement of interfoliar buds may be due to a group effect, where each developing DS inhibits the more proximal DSs in the long shoot terminal bud.     

莲的根茎构造,伸长与增粗

莲 (Nelumbo nucifera)种苗的茎短而直立,叶互生。幼苗期茎延伸成横卧根茎,其上生有营养芽及混合芽。腋生鳞片为叶的衍生部分,形如叶鞘状,包着预芽。年苗上的冬芽内全为混合芽。根茎内的维管束分散排列,无导管及形成层存在。节间延长通过肋状分生组织及节间内的薄壁组织细胞分裂与增长来完成。根茎可由初生加厚分生组织,维管束细胞,皮层薄壁细胞等的细胞分裂,使层次增加,但增粗主要是由皮层薄壁细胞体积显著增大而引起的。     

ORGANOGRAPHY,BRANCHING, AND THE PROBLEM OF LEAF VERSUS BUD DIFFERENTIATION IN THE VINING EPIPHYTIC FERN GENUS MICROGRAMMA

Investigation of the development and organography of the shoot systems of Microgramma vacciniifolia and M. squamulosa was undertaken for the purpose of determining: (1) the features of shoot growth that are responsible for the distinctive vining character of these epiphytic ferns; and (2) the mode of origin of branches and their contrast with leaf initiation. Shoots of both species are dorsiventral and plagiotropic (i.e., parallel to the substrate) in habit. Since the shoot apical meristem is radial in transectional symmetry, shoot dorsiventrality in Microgramma is a postgenital or secondary developmental event, and its inception is related to the initiation of lateral appendages. Leaves and buds arise in a distichous phyllotaxis and occupy opposite and alternating positions on the dorsal surfaces and flanks of the rhizome. Endogenous roots are initiated in two rows from the ventral surface of the stem, in the vicinity of the rhizome meristem; however, they do not emerge from the rhizome until some distance behind the tip and do not elongate until the region of substrate contact. We conclude that the vining nature of this fern rhizome is a result of precocious internodal elongation and the concomitant delay of leaf and bud expansion in the region of stem elongation. In addition, observation of branch origin confirms previous suggestions that branching in Microgramma is strictly lateral and extra-axillary and not a dichotomous derivative as proposed by some workers. Leaf and bud primordia differ not only in the nature of their respective vascular supplies but also in their actual course of initiation. In the case of the leaf, the primordium is precociously emergent and exhibits a lenticular apical cell at its summit when it is only one plastochron removed from the flanks of the apical meristem. By contrast, initials of the bud primordium divide less actively and remain in a sunken position for at least 5–6 plastochrons; only when the bud apex becomes expanded and emergent does a tetrahedral apical cell become recognizable at the tip of the bud promeristem. Because of the distinctive pattern of branch and leaf origin, as well as the lack of adventitious and phyllogenous origin of branch primordia, we suggest that the shoot of Microgramma is a useful test organism for the re-examination of the problem of leaf and bud determination in the ferns.     

Organ correlations and flowering in chenopodium rubrum L.

Correlations within a shoot ofChenopodium rubrum L. ecotype 374 grown under continuous light or photoperiodic flower induction were studied using surgical treatments. Removal of a single pair of shoot organs had a variety of effects depending on position: significant changes in the number of leaf pair on the main axis or in axillary buds and in the height of shoot apices; or no effect on the parameters scored. Flowering was not affected by any of the treatments carried out. Decapitation brought about a significant increase in the number of leaf pairs in axillary buds and flowering was inhibited in 8- and 9-d old plants. Flowering was not affected in 21-d old plants. The role of shoot organ correlations, especially that of apical dominance, in regulation of flowering inC.rubrum is discussed.     

Phragmites die-back: toxic effects of propionic, butyric and caproic acids in relation to pH

Symptoms which are associated with die-back in Phragmites : growth inhibition, root and bud death, premature shoot senescence, blocked aeration and vascular systems, especially in rhizomes and roots, and abnormal surface and internal cell-wall lignification and suberization of roots were induced by each of three of the lower volatile organic acids, propionic, butyric and caproic. These acids were applied in nutrient media in concentrations similar to those previously associated with die-back sites and/or in sediments containing rotting rhizomes and roots of the plant. At concentrations of 1.4 and 0.56 mM, respectively, butyric and caproic acids were each found to be highly toxic at pH 4.5, but relatively innocuous at pH 6. Propionic acid, applied at a much higher concentration of 10.4 mM, was highly toxic at both pH 4.5 and 6. The results support previous findings that the undissociated forms of the organic acids are the more toxic. Rhizomes and roots, rotting in water or waterlogged sand, released cocktails of acids and produced pH in the range 4.8–5.4. Phragmites seedlings planted in these media died within 12 h. Overall, the results support the theory that die-back in Phragmites can be induced and/or perpetuated by organic acids released from the decaying underground parts of the plant or other sources of organic matter.     

Hormone Levels and Apical Dominance in the Aquatic Fern Marsilea drummondii A. Br

Terminal buds and successively subjacent lateral buds of the water fern, Marsilea drummondii, were examined to determine the pattern of hormone distribution in relation to apical dominance. Quantitative levels of indole-3-acetic acid (IAA), abscisic acid (ABA), zeatin and zeatin riboside (Z and ZR), and isopentenyladenosine (iPA) were determined by a solid-phase immunoassay using polycional antihormone antibodies. Enzyme-linked immunosorbent assay was used following a one-step HPLC purification procedure to obtain the free hormones. Active shoot apices contained the most IAA and Z-type cytokinins and inhibited buds the least. No significant differences in ABA levels were found leading to the conclusion that ABA did not play any role in apical dominance. The normal precedence of the most rapid outgrowth of the youngest inhibited bud as observed previously in decapitated plants was well correlated with its very high level of iPA observed in this study. The same phenomenon was observed in the median buds but with a weaker amplitude. The presence of this storage form could indicate that a bud at its entry into quiescence eventually looses the ability to hydroxylate iPA to Z-type cytokinins when it is fully inhibited. IAA and Z + ZR are concluded to be essential for lateral bud growth.     

High Frequency Shoot Organogenesis in Sorghum bicolor (L) Moench

In vitro morphogenesis that includes enlargement of apical meristems and differentiation of shoot buds on the surface of enlarged meristemoids, have been observed in Sorghum bicolor. The enlargement of apical meristems occurred on the MS medium containing different auxins [2,4-dichlorophenoxyacetic acid (2,4-D), 2,4,5-trichloro-phenoxyacetic acid (2,4,5-T) and parachlorophenoxyacetic acid (pCPA)] with or without 6-benzylaminopurine (BAP)]. Large number of shoot buds arose over the entire surface of enlarged, green, compact, nodular, hard and shiny meristemoids on transfer to medium containing BAP (2.0 mg l^?1) + (IAA) indole 3-acetic acid (0.5 mg l^-1). Histological observations revealed the origin of shoot apices from the surface of enlarged meristemoids. Efficient rooting was achieved onto half-strength MS medium supplemented with α-napthaleneacetic acid (NAA, 2.0 mg l^?1) and sucrose 2% (w/v). Plantlets were transferred to earthen pots under field conditions for the evalution of several agronomically important characters.     

SYMMETRY AND DEVELOPMENT OF BUTOMUS UMBELLATUS (BUTOMACEAE) AND LIMNOCHARIS FLAVA (LIMNOCHARITACEAE)

The prostrate rhizome of Butomus umbellatus produces branch primordia of two sorts, inflorescence primordia and nonprecocious vegetative lateral buds. The inflorescence primordia form precociously by the bifurcation of the apical meristem of the rhizome, whereas the non-precocious vegetative buds are formed away from the apical meristem. The rhizome normally produces a branch in the axial of each foliage leaf. However, it is unclear whether the rhizome is a monopodial or a sympodial structure. Lateral buds are produced on the inflorescence of B. umbellatus either by the bifurcation or trifurcation of apical meristems. The inflorescence consists of monochasial units as well as units of greater complexity, and certain of the flower buds lack subtending bracts. The upright vegetative axis of Limnocharis flava has sympodial growth and produces evicted branch primordia solely by meristematic bifurcation. Only certain leaves of the axis are associated with evicted branch primordia and each such primordium gives rise to an inflorescence. The flowers of L. flava are borne in a cincinnus and, although the inflorescence is simpler than that of Butomus umbellatus, the two inflorescences appear to conform to a fundamental body plan. The ultimate bud on the inflorescence of Limnocharis flava always forms a vegetative shoot, and the inflorescence may also produce supernumerary vegetative buds. Butomus umbellatus and Limnocharis flava exhibit a high degree of mirror image symmetry.     

Callus, Organogenesis and Plantlet Formation in Tissue Cultures of Clitoria ternatea

Decoated seeds of Clitoria ternatea L. germinated on Murashigeand Skoog (Physiologia Plantarum 1962, 15, 473–97) basalmedium (BM) and differentiated callus and bipolar embryoids(two-step method) in low frequency. Calluses developed on lateralroots [BM+KN(0.1 mg 1^–1)], on roots and hypocotyls [BM+KN(0.5mg 1^–1)], and on roots [BM+KN+IAA (0.5 mg 1^–1 ofeach)]. On basal medium with KN (0.5 mg 1^–1) and withKN+IAA (0.5 mg1^–1 of each), multiple shoot buds and embryoids(one-step method) were differentiated directly on split hypocotylsand roots. In the former, shoot buds developed even on unsplithypocotyls. Rhizogenesis on isolated shoot buds occurred efficientlyin BM+indole butyric acid (IBA 0.1 mg 1^–1) and BM+IAA(0.1 mg 1^–1 and 0.5 mg 1^–1). Profuse direct embryoidsand shoot buds developing on root systems are interesting morphogeneticphenomena rarely reported. Clitoria ternatea L., callus, embryoids, multiple shoot buds, regeneration     

The maryland mammoth allele and rooting both perturb the fate of florally determined apices in Nicotiana tabacum.

The stability of the florally determined state in terminal and axillary buds of two tobacco cultivars was studied. We used Hicks and Hicks Maryland Mammoth, near-isogenic cultivars of Nicotiana tabacum differing at the recessive maryland mammoth locus which confers short-day behavior. The experimental design consisted of growing plants in short-day conditions and subjecting them to three bioassays in long-day conditions: in vitro culture of apices consisting of meristems and three to four leaf primordia; rooting of buds consisting of meristems and 8 to 12 leaves, leaf primordia, and internodes; and release from apical dominance of axillary buds in situ. Cultured terminal and axillary apices expressed floral determination, indicating that meristems can be florally determined. Two lines of evidence indicate that rooting destabilizes an already acquired florally determined state: cultured apices from both axillary and terminal buds produced fewer nodes after excision than homologous buds which were rooted; and a lower percentage of rooted axillary buds from Hicks Maryland Mammoth plants expressed floral determination than did homologous axillary buds grown out in situ in noninductive conditions. Rooted buds from the two genotypes expressed floral determination at different frequencies, but produced abnormal inflorescences at similar frequencies, indicating that roots and the maryland mammoth allele influence common as well as unique processes associated with floral determination.     

The Combined Effects of Salinity and Root Anoxia on Growth and Net Na+ and K+-accumulation in Zea mays Grown in Solution Culture

The effects of excess salinity and oxygen deficiency on growthand solute relations in Zea mays L. cv. Pioneer 3906 were examinedin greenhouse experiments. The roots of plants 14 d old growingin nutrient solution containing additions of NaCl in the range1.0–200 mol m^–3 were either exposed to a severedeficiency of O₂ by bubbling with nitrogen gas (N₂ treatment),or maintained with a supply of air (controls), for a periodof 1–7 d. The threshold NaCl concentration resulting inappreciable inhibition of leaf extension, and shoot f. wt gainin controls was between 10 and 25 mol m^–3. At 25 mol m^–3NaCl the ratio of Na+/K+ transported to shoots was about 20times greater than in plants in 1.0 mol m^–3 NaCl. Theeffect of addition of NaCl to the nutrient solution was to enhanceNa+ movement but simultaneously depress the rate of K+ transportto shoots (per g f. wt roots). Interactions between NaCl levels and aeration treatment wereshown by analyses of variance to be statistically significantfor leaf extension, shoot and root f. wt gains, Na+ and K+ concentrationsin shoots and roots. When roots were N₂-treated, shoot and rootgrowth were depressed, the effect of aeration treatment beinggreatest at NaCl concentrations of 50 mol m^–3 or less.Additionally, N₂-treatment greatly accelerated Na- transportto shoots while depressing K+ transport still further, so thatat 10 mol m^–3 NaCl the ratio Na+/K+ acquired by the shootswas 230 times greater than in controls. Over the concentrationrange 1.0 to 50 mol m^–3 NaCl, the ratio Na+/K+ transportedto shoots by anoxic roots increased by a factor of 860. Mechanisms controlling changes in solute flux to the shoot,and the significance in relation to plant tolerance of excesssalts or oxygen deficiency are discussed. Anaerobic, corn, flooding, maize, oxygen-deficiency, salinity