RELATIONSHIPS BETWEEN FLOODING TOLERANCE,LIFE HISTORY,AND SHORT-TERM COMPETITIVE PERFORMANCE IN THREE SPECIES OF POLYGONUM

The objective of this study was to compare the growth and short-term (single season) competitive performance of three species of Polygonum known to differ in flooding tolerance and life history. Polygonum amphibium is a perennial with low sexual reproductive effort and a relatively high degree of flooding tolerance, P. lapathifolium is an annual species with a high sexual reproductive effort and a low tolerance to flooding, and P. hydropiperoides is intermediate to the other two in terms of sexual reproductive effort and flooding tolerance. In order to determine the relative growth and competitive abilities of these species, mixtures and monocultures of plants were grown in pots and maintained under three flooding regimes: 1) flooded, 2) partially drained, and 3) well drained. Both P. hydropiperoides and P. amphibium grew best under flooded and partially drained conditions with reduced growth in the drained treatment. Polygonum lapathifolium, in contrast, grew as well in the drained treatment as in the more flooded treatments. Results from competition experiments were consistent in showing the relative competitive abilities to be P. lapathifolium > P. hydropiperoides > P. amphibium regardless of flooding regime. Thus, short-term competitive performance was found to trade off with flood tolerance rather than with sexual reproductive effort.     

两栖蓼的分子系统学研究

两栖蓼是一种水陆两栖植物,植株在不同生态环境下外部形态差异较大,同时两栖蓼的系统位置存在争议,被归入春蓼组(sect.Persicaria)或提升为两栖蓼组(sect.Amphibium)。该文选取两栖蓼及春蓼组植物12种,以及刺蓼组、头状蓼组、神血宁组、拳参组、萹蓄组和外类群掌叶大黄共23种植物进行研究。植物总DNA的提取采用改进的CTAB法,所测序列以及从Genbank数据库下载的序列,以掌叶大黄为外类群,采用最大简约法和贝叶斯法对核糖体ITS序列和叶绿体trn L-F序列进行了系统发育分析。ITS序列对位排列的长度为735 bp,包括489个可变位点,272个位点是信息位点。简约法得到9个简约树,步长为1 084,CI指数为0.680,RI指数为0.614。trn L-F序列对位排列的长度为1 121 bp,包括427个可变位点,239个位点是信息位点。简约法寻找到9个简约树,步长为551(CI=0.911,RI=0.910)。贝叶斯法和简约法得到的树基本一致。分子序列分析结果显示,trn L-F序列树类似于ITS序列树。ITS序列构建的发育树上,两栖蓼与刺蓼组植物、春蓼组其他植物形成3个并列的分支;在trn L-F序列树上,两栖蓼则与其他春蓼组植物形成两个并列的分支。由此可见,两栖蓼与春蓼组其他植物的亲缘关系较远,成一独立的分支。两个分子证据支持将两栖蓼提升为两栖蓼组的处理意见。此外,两栖蓼的花粉具散沟,与典型的春蓼组的具散孔花粉不一致。再加上两栖蓼水陆两栖的特性,因此支持把两栖蓼提升为两栖蓼组的观点。两栖蓼组的界定为多年生草本,水陆两栖,根状茎横生,生于水中茎漂浮,叶长圆形或椭圆形,生于陆地茎直立,叶披针形或长圆状披针形,托叶鞘为筒状、薄膜质,总状花序穗状,瘦果近圆形,花粉具散沟。     

GEITLERINEMA SPECIES (OSCILLATORIALES,CYANOBACTERIA) REVEALED BY CELLULAR MORPHOLOGY,ULTRASTRUCTURE, AND DNA SEQUENCING1

Geitlerinema amphibium (C. Agardh ex Gomont) Anagn. and G. unigranulatum (Rama N. Singh) Komárek et M. T. P. Azevedo are morphologically close species with characteristics frequently overlapping. Ten strains of Geitlerinema (six of G. amphibium and four of G. unigranulatum) were analyzed by DNA sequencing and transmission electronic and optical microscopy. Among the investigated strains, the two species were not separated with respect to cellular dimensions, and cellular width was the most varying characteristic. The number and localization of granules, as well as other ultrastructural characteristics, did not provide a means to discriminate between the two species. The two species were not separated either by geography or environment. These results were further corroborated by the analysis of the cpcB‐cpcA intergenic spacer (PC‐IGS) sequences. Given the fact that morphology is very uniform, plus the coexistence of these populations in the same habitat, it would be nearly impossible to distinguish between them in nature. On the other hand, two of the analyzed strains were distinct from all others based on the PC‐IGS sequences, in spite of their morphological similarity. PC‐IGS sequences indicate that these two strains could be a different species of Geitlerinema. Using morphology, cell ultrastructure, and PC‐IGS sequences, it is not possible to distinguish G. amphibium and G. unigranulatum. Therefore, they should be treated as one species, G. unigranulatum as a synonym of G. amphibium.     

Differences in morphology and reproductive traits of Galerucella nymphaeae from four host plant species

The water lily beetle Galerucella nymphaeae L. (Coleoptera: Chrysomelidae) exploits different hosts, including Nuphar lutea Sm. and Nymphaea alba L. (both Nymphaeaceae), as well as Polygonum amphibium L. and Rumex hydrolapathum Hudson (both Polygonaceae). The present study investigates whether within-species differences in morphological and reproductive traits are associated with differences in host species exploitation. A total of 1103 adult beetles were collected from 11 localities in The Netherlands, one of which contained all four hosts and three other localities contained hosts from both families (sympatric localities). Adults originating from Nuphar and Nymphaea were on average darker in colour and larger in size and had disproportionally bigger mandibles than beetles originating from Polygonum and Rumex across the 11 localities. Head capsules of first instar larvae from Nymphaeaceae hosts were between 17% and 28% larger than those of larvae from Polygonaceae hosts. Furthermore, beetles from Nuphar and Nymphaea laid larger sized eggs, but fewer eggs per clutch than beetles originating from Polygonum and Rumex. Although host related variation was less pronounced at the sympatric localities than in the allopatric localities, differences in larval and adult size were still highly significant at the sympatric localities. It is not clear whether the observed differences are genetically based, as opposed to host induced. However, leaf toughness varied among species in a way suggesting that leaf toughness may be partly responsible for host related differences in G. nymphaeae.     

三种水库消落带草本植物对完全水淹的适应机制研究

为了解虉草(Phalaris arundinacea)、牛鞭草(Hemarthria altissima)和狗牙根(Cynodon dactylon)3种水库消落带草本植物在完全水淹条件下的生理生态适应机制,对这3种植物的生态指标(枝条、叶和生物量)和生理指标(可溶性糖、淀粉和根系活力)的动态变化特征进行了研究。结果表明：在完全水淹条件下,虉草、牛鞭草和狗牙根都不产生新的枝条,它们通过减少总叶数、总叶长和生物量的方式来适应水淹环境。虉草和狗牙根通过减缓枝条生长速率来适应水淹环境,而牛鞭草则是通过先加快生长后抑制生长来保存活力。虉草、牛鞭草和狗牙根均以少量的碳水化合物(可溶性糖和淀粉)消耗,降低根系活力等方式来适应长期的水淹环境。3种草本植物的水淹耐受能力由大到小依次为狗牙根、虉草、牛鞭草。     

Heterophylly results in a variety of “spectral signatures” in aquatic plant species

The leaf reflectance spectra (280–887 nm) of two heterophyllous aquatic plant species Polygonum amphibium (L.) and Nuphar luteum (L.) were compared and their relation to physical properties of the leaves examined. In P. amphibium contrasting environmental conditions along water–land gradient affected the majority of anatomical and morphological properties of leaves, but less differences were observed in photosynthetic pigment and total flavonoid contents. Leaf mass per area (LMA), palisade mesophyll, leaf thickness, trichome length and anthocyanin content per dry mass were correlated to the different parts of spectra. In N. luteum natant and submerged leaves differed significantly in all measured parameters. Chlorophyll a, anthocyanin and carotenoid contents per dry mass were related to reflectance in the red region, while leaf thickness, anthocyanin and total flavonoid contents per leaf area were related to reflectance in the near infrared region. Redundancy Analysis (RDA) indicated that in P. amphibium the average length of trichomes and LMA explained 72% and 6% variability of the spectra, whereas in N. luteum anthocyanin content per dry mass, explained 57% variability of the spectra. The comparison of natant leaves of both species showed that they were more similar than different leaf types within the single species.     

Relative effects of Justicia americana litter on germination,seedlings and established plants of Polygonum lapathifolium

Justicia americana (L.) Vahl is one of the most common species associated with seasonally variable aquatic habitats in the south-central U.S.A. When water levels decline, litter of Justicia decomposes on the exposed mudflats where most emergent aquatic species typically germinate and establish. In this study, we have tested the effects of aqueous leachates of Justicia shoots on the germination, seedling growth and established plant growth of Polygonum lapathifolium L., a common annual colonizer of exposed mudflats. Maximal leachate concentrations were prepared using quantities of litter comparable to those estimated for natural stands of Justicia. At these concentrations, seed germination and transplant growth were unaffected, but seedling growth was inhibited. When Polygonum was tested with lower concentrations of Justicia leachate, germination was still unaffected, but seedling growth was found to be enhanced, especially at the lowest concentrations. Using mannitol as an inert osmotic agent, Polygonum seedling growth was found to be inhibited at osmotic potentials equivalent to those of all leachate concentrations. Thus, the effects of leachate on seedling growth include both osmotic and non-osmotic components. Overall, the results support the hypothesis that the seedling stage of the plant life cycle is the stage most susceptible to permanent effects by the chemicals released from decomposing litter.     

Benthic macroinvertebrates associated with four species of macrophytes

Benthic macroinvertebrates associated with four species of macrophytes (Nymphoides peltata, Ceratophyllum demersum, Polygonum amphibium and Carex sp.) were investigated during two growing seasons (2001 and 2002) in the slow-flowing Čonakut Channel in the Kopački rit Nature Park in Croatia. A total of 31 macroinvertebrate taxa were found. C. demersum, a submerged plant with dissected leaves, supported the highest macroinvertebrate abundance, almost seven times more than N. peltata, a floating plant with undissected leaves, which harboured the lowest abundance during the research period. Chironomidae larvae (50–83%) and Oligochaeta (14–46%) were the most abundant groups recorded on all macrophyte species. Water-level fluctuation, because of its influence on the appearance and growth of aquatic vegetation, and the trophic state of water within the macrophyte stands seemed to be the main factors which affected the taxonomic composition and abundance of macroinvertebrates.     

Ethnobotany of folk medicinal aquatic plants in Jordan

Eighty-seven species belonging to 59 genera and 33 plant families identified in the study area are presented. The three families with the most species represented were Labiatae (nine aquatic species), Compositae (seven species), and Salicaceae (seven species). The genera most represented wereMentha (six species),Polygonum (five species), andSalix (five species). Sixty-three folk-medicinal aquatic species (73.3%) had similar therapeutic uses in neighboring countries, while the 24 remaining species (26.7%) did not show therapeutic similarity with their use in other countries. Emerged species (plants rooted in soil under water but which emerge partially above the water’s surface) were the most recorded, while amphibious, submerged, and floating species were the least recorded. The folk-medicinal importance of the recorded aquatic species were classified by rank-order priority (ROP). Twenty-one species (24%) had ROP values higher than 50, indicating the highest popularity level in folk-medicinal potentiality; 26 species (29.9%) had therapeutic effects informed by fewer than three informants and were therefore excluded from further consideration; 40 species (46.1%) had ROP values of less than 50, and were thus classified as nonpopular medicinal plants.     

Effect of complete plant submergence at different growth stages on grain yield,yield components and nutrient content of rice

Summary Of the three growth stagesviz., seedling establishment, maximum tillering and flowering, complete submergence of plant at flowering stage was found to be most critical followed by seedling estabilishment and maximum tillering stages. Among the three stages of reproductive growth phase, booting stage was found to be most susceptible to complete submergence followed by flowering and post-flowering. The submergence at booting for 4 days was equally detrimental as that of 6 or 8 days at flowering. Irrespective of the growth stages, the plants subjected to complete submergence showed higher nitrogen content (in plant as well as in grain) as compared to those grown under control conditions (5±2 cm) and increased with the increase in duration of submergence. The P and K contents in the plant decreased under submergence.     

Morphological and anatomical changes of Melaleuca cajuputi under submergence

Melaleuca cajuputi is a woody plant of the Myrtaceae which is a dominant species in tropical peat swamps in southern Thailand, where the groundwater level fluctuates greatly. Although the current year seedlings are likely submerged, their adaptive responses have never been studied. The objective of the present study was to examine their responses to submergence, and especially their morphological and anatomical changes. Not only did the seedlings of M. cajuputi survive submergence for 56 days, but they could also increase their dry weight, shoot length, and leaf number during submergence. These growth responses to submergence indicate that the seedlings of M. cajuputi could make photosynthetic production under water. The leaves that developed under water were heterophyllous “aquatic leaves” that appear to represent adaptations to improve the uptake of gases from the water. Intercellular spaces in the stems and leaves were more strongly developed in the submerged seedlings than in non-submerged seedlings with the shoot and leaves in the air. The intercellular spaces appear to be schizogenous aerenchyma that facilitates gas exchange. The growth responses and anatomical responses in stems and leaves to submergence, which were found in M. cajuputi, are commonly known in herbaceous plants with amphibious characteristics, but had not been reported in woody plants. And our results suggest that M. cajuputi adapts to submergence similarly to other amphibious plants, thereby ensuring continuing biomass production.     

Flavonoid chemistry ofPolygonum sect.Tovara (Polygonaceae): A systematic survey

Polygonum sect.Tovara includes three controversial species;P. virginianum, P. filiforme, andP. neofiliforme. The flavonoid chemistry of these was examined to provide additional information on their delimitation and levels of differentiation. Eight flavonoid compounds were isolated and identified, all of which were 3-O-glycosides of the flavonols kaempferol, quercetin, and myricetin, and their acylated derivatives. Although they exhibit relatively simple flavonoid profiles, the three taxa are readily distinguished by their flavonoid constituents. In addition, they show fundamental differences in flavonol types and glycosylation patterns. These results, in conjunction with evidence from the morphology, strongly suggest thatP. virginianum, P. filiforme, andP. neofiliforme are closely allied but distinct species.     

Metal content in higher aquatic plants in a small siberian water reservoir

The dynamics of metal content in higher aquatic plants (macrophytes) in a small Bugach water reservoir in 1998–2006 was studied. A comparative estimation of the metal content in six macrophyte species (Typha latifolia L., Typha angustifolia L., Polygonium amphibium L., Potamogeton perfoliatus L., Potamogeton pectinatus L., Phragmites australis (Cav) Trin. Ex Steud.) showed that their metal concentrations do not generally exceed those known from the literature. Cluster analysis showed that the macrophyte species under study form two ecological groups with respect to the metal content, i.e., submerged plants (hydrophytes) and emergent aquatic plants (heliophytes).     

水体溶氧影响陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)对完全水淹的耐受力

溶氧是水环境中一个重要的环境因子,为了探讨水中的溶氧含量水平是否会对陆生植物的耐淹能力造成影响,研究了陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)在遭受不同溶氧含量水体完全淹没后的生长表现、存活情况和非结构碳水化合物的变化。实验结果表明:(1)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的存活。受高溶氧水体完全水淹的喜旱莲子草和牛鞭草主茎的完好程度和存活叶的数量均显著高于遭受低溶氧水体完全水淹的喜旱莲子草和牛鞭草,喜旱莲子草和牛鞭草在高溶氧水体完全水淹后的生物量比低溶氧水体完全水淹后要高;(2)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的生长,受高溶氧水体完全水淹的喜旱莲子草主茎伸长生长和不定根生长显著强于受低溶氧水体完全水淹的喜旱莲子草,在不定根的生长上牛鞭草也具有同样的表现。(3)高溶氧水环境有利于减小被完全淹没的喜旱莲子草和牛鞭草的碳水化合物消耗,两种植物在受高溶氧完全水淹后体内具有的非结构性碳水化合物含量均比受低溶氧完全水淹后高。(4)喜旱莲子草比牛鞭草能更好地耐受完全水淹,当处于低溶氧完全水淹时表现得更为明显,本研究表明入侵物种喜旱莲子草比本地物种牛鞭草具有更强的环境适应能力和水淹耐受能力。     

Hypoxic Quiescence in Alligatorweed

Propagules (stem cuttings) of alligatorweed [Alternanthera philoxeroides (Mart.) Griseb.] were subjected to various treatments of submergence, emergence, light, dark, and oxygen to relate hypoxia to the suppression of alligatorweed growth by dark submergence. In the dark, submergence completely inhibited alligatorweed growth. A nitrogen atmosphere inhibited growth from emersed propagules in the dark. The suspension of growth by dark submergence was relieved by light (40 μ einsteins/m²× s photo-synthetically active radiation), emergence (whereby air is transported to underwater nodes via the aerenchyma), or exposure to 100 mg · l^-1 of hydrogen peroxide solution renewed at 48-h intervals. Suspension of growth by a nitrogen atmosphere was relieved by inclusion of 16% oxygen in the ambient atmosphere. Dark to light transfers showed that the suspension of growth was immediately relieved if the requisite conditions were restored; this phenomenon, as described, thus appears to be hypoxic quiescence. The suppression of growth of two other amphibious species, mild smartweed (Polygonum hydropiperoides Michx.) and creeping water-primrose (Jussiaea repens L. var. glabrescens Ktze.), by dark submergence suggests that hypoxic quiescence may be a factor in the life history of amphibious species other than alligatorweed.     

Aquatic adventitious root development in partially and completely submerged wetland plants Cotula coronopifolia and Meionectes brownii

Background and Aims

A common response of wetland plants to flooding is the formation of aquatic adventitious roots. Observations of aquatic root growth are widespread; however, controlled studies of aquatic roots of terrestrial herbaceous species are scarce. Submergence tolerance and aquatic root growth and physiology were evaluated in two herbaceous, perennial wetland species Cotula coronopifolia and Meionectes brownii.

Methods

Plants were raised in large pots with ‘sediment’ roots in nutrient solution and then placed into individual tanks and shoots were left in air or submerged (completely or partially). The effects on growth of aquatic root removal, and of light availability to submerged plant organs, were evaluated. Responses of aquatic root porosity, chlorophyll and underwater photosynthesis, were studied.

Key Results

Both species tolerated 4 weeks of complete or partial submergence. Extensive, photosynthetically active, aquatic adventitious roots grew from submerged stems and contributed up to 90 % of the total root dry mass. When aquatic roots were pruned, completely submerged plants grew less and had lower stem and leaf chlorophyll a, as compared with controls with intact roots. Roots exposed to the lowest PAR (daily mean 4·7 ± 2·4 µmol m⁻² s⁻¹) under water contained less chlorophyll, but there was no difference in aquatic root biomass after 4 weeks, regardless of light availability in the water column (high PAR was available to all emergent shoots).

Conclusions

Both M. brownii and C. coronopifolia responded to submergence with growth of aquatic adventitious roots, which essentially replaced the existing sediment root system. These aquatic roots contained chlorophyll and were photosynthetically active. Removal of aquatic roots had negative effects on plant growth during partial and complete submergence.     

鄱阳湖洲滩湿地植物生长和营养繁殖对水淹时长的响应

水淹时长是影响湿地植物分布的重要因素。在水情不断变化的背景下,鄱阳湖洲滩湿地植物种群和群落的变化规律还不清楚。为了探究淹水时长对鄱阳湖洲滩3种优势植物生长和繁殖的影响,并预测在水文发生剧烈变化后,3种优势植物分布的趋势,采用控制实验模拟了不同水淹时长(0、60、90、120、150 d和180 d)下鄱阳湖湿地3种优势植物(灰化薹草(Carex cinerascens)、南荻(Miscanthus lutarioriparius)和虉草(Phalaris arundinacea))的生长和繁殖情况。研究结果表明:1)南荻在水淹超过120 d后,存活率开始降低,水淹到达180 d完全死亡。而灰化薹草和虉草在淹水180 d后仍全部存活。表明南荻耐淹水的能力弱于其他两种植物。2)水淹显著降低灰化薹草的总生物量,并且总生物量随水淹时长的增加而逐渐降低。而短时间(小于150 d)的水淹没有对虉草总生物量产生显著影响。3)退水初期,灰化薹草的恢复生长趋向于地上部分,而虉草表现为地下和地下部分同步生长。该研究结果可以为预测水文情势变化下鄱阳湖湿地植物种群和群落的动态变化提供依据。     

Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae)

The pollen of 30 taxa (27 species, one subspecies and two varieties) in two genera, viz Polygonum s. str. and Polygonella was investigated with LM and SEM, and some selected taxa with TEM. In all genera investigated the pollen is prolate to spheroidal, and the aperture is mostly tricolporate, rarely panto-hexacolporate (especially Polygonum section Polygonum). The exine sculpturing pattern is the most variable feature. Three types of exine can be recognized. Type 1 (Avicularia-Type, sensu Hedberg) - All species of section Polygonum and section Tephis share the smooth tectate exine with spinules, sometimes the surface is more or less rough (Polygonum afromontanum in section Tephis). Type 2 (Pseudomollia-Type, sensu Hong) - Pollen of Polygonum molliaeforme (section Pseudomollia) has the exine, which is verrucose on both poles and nearby the mesocolpium, and mostly psilate around the ectoaperture. Type 3 (Duravia-Type, sensu Hedberg) - Pollen grains of Polygonum section Duravia and Polygonella have the exine which is semitectate-reticulate at the mesocolpium and the poles, and rugulate/reticulate or sometimes foveolate with microspinules around the ectoapertures. The pollen grains in four taxa (viz Polygonum section Pseudomollia, P. section Duravia and genus Polygonella) have a well-marked dimorphism of the ektexine, which is considered to be a synapomorphic condition. The differences of pollen grain between the genus Polygonella and Polygonum section Duravia are almost non existent and clearly interrelated. It is therefore postulated that the similarity in pollen of both taxa is not the result of convergency, but is interpreted as a homology. It is noteworthy that the pollen of Polygonum molliaeforme (section Pseudomollia) appears as intermediate between the Avicularia-type and the Duravia-type, and is well supported the value of separated section for its own. Additionally, in TEM, some exine ultrastructures (e.g. columellae, foot layer, endexine) appear to be valuable characters for comparison between/among taxa. The systematic potentialities of the pollen data of the studied taxa at various systematic levels are also discussed.     

Light acclimation, CO2 response and long-term capacity of underwater photosynthesis in three terrestrial plant species

To characterize underwater photosynthetic performance in some terrestrial plants, we determined (i) underwater light acclimation (ii) underwater photosynthetic response to dissolved CO₂, and (iii) underwater photosynthetic capacity during prolonged submergence in three species that differ in submergence tolerance: Phalaris arundinacea, Rumex crispus (both submergence-tolerant) and Arrhenatherum elatius (submergence-intolerant). None of the species had adjusted to low irradiance after 1 week of submergence. Under non-submerged (control) conditions, only R. crispus displayed shade acclimation. Submergence increased the apparent quantum yield in this species, presumably because of the enhanced CO₂ affinity of the elongated leaves. In control plants of the grass species P. arundinacea and A. elatius, CO₂ affinities were higher than for R. crispus. The underwater photosynthetic capacity of R. crispus increased during 1 month of submergence. In P. arundinacea photosynthesis remained constant during 1 month of submergence at normal irradiance; at low irradiance a reduction in photosynthetic capacity was observed after 2 weeks, although there was no tissue degeneration. In contrast, underwater photosynthesis of the submergence-intolerant species A. elatius collapsed rapidly under both irradiances, and this was accompanied by leaf decay. To describe photosynthesis versus irradiance curves, four models were evaluated. The hyperbolic tangent produced the best goodness-of-fit, whereas the rectangular hyperbola (Michaelis-Menten model) gave relatively poor results.