Assessment of aquatic and terrestrial reed (Phragmites australis) stands

A survey of recent publications shows that research on Phragmites australishas oftenapplied character because of the considerableecological and economic significance of the species.The main applications are water treatment, agriculture(food production or weed control) and natureconservation. In Europe, most research on natural reedstands has been motivated by reed die-back and effortstowards protection or restoration. Reed progressionand reed control have been the main concerns in otherparts of the world, and reed progression has alsoreceived increasing attention in Europe. While reeddie-back generally affects aquatic stands, progressioncan occur at both terrestrial and aquatic sites, andit can be desired (e.g. lake shore restoration)or unwanted (e.g. in species-rich fens ormarshes). Therefore, reed stands need to be assessedindividually to decide on management aims andappropriate methods. The varying status of Phragmites australisformed the background of the`European Reed Conference' held in Zürich/Switzerlandin October 1998. The seven contributions published inthis special issue are introduced with particularreference to differences between aquatic andterrestrial reed stands and to approaches used in theirassessment.     

Strategies of nitrogen cycling of Phragmites australis at two sites differing in nutrient availability

Below-ground biomass and nitrogen content were determined at two genetically homogeneous Phragmites stands differing in morphology, in productivity, and in nutrient supply. Comparable ratios between above-ground standing crop and rhizome biomass were found at both sites, whereas the root biomass/above standing crop ratio was significantly higher at the nutrient poor site. Investigations on the dynamics of nitrogen content revealed distinct differences in nitrogen translocation to the rhizomes between the investigated clones indicating two ecophysiological strategies in storage behaviour. These two strategies could be attached to the “assimilation type” and to the “translocation type”, respectively. A modified definition of both types is presented.     

互花米草与芦苇光合色素含量对淹水措施的响应

以上海崇明东滩入侵植物互花米草与本地种芦苇为研究对象,研究持续淹水胁迫对两种植物光合色素含量的影响.结果表明：互花米草与芦苇叶片的光合色素基本组成与含量不同,对持续淹水的响应也不同.在持续淹水胁迫下,互花米草叶绿素a、叶绿素b及类胡萝卜素含量降低,叶绿素a／b、类胡萝卜素／叶绿素值提高;芦苇各色素含量升高,叶绿素a／b、类胡萝卜素/叶绿素值基本保持稳定.解除持续淹水胁迫后,互花米草各色素含量逐渐升高,叶绿素a／b、类胡萝卜素／叶绿素值降低,并逐渐接近对照水平;而芦苇各色素含量显著高于对照.两种植物均表现出一定的补偿效应,但芦苇比互花米草更能适应同等程度的持续淹水胁迫.应用持续淹水措施治理互花米草时,可采用本地种芦苇作为治理后湿地恢复的替代植物.     

Differences in photorespiration,glutamine synthetase and polyamines between fragmented and closed stands of Phragmites australis

Physiological processes related to C and N metabolism were investigated in closed healthy, and fragmented die-back stands of Phragmites australis (Cav.) Trin. ex Steudel along the shores of Lake Balaton, Hungary. In the leaves, similar concentrations of total N and P, K⁺, Na⁺, Ca²⁺ and Mg²⁺ were found. However, higher concentrations of soluble proteins in the fragmented stand indicated alterations in N metabolism. In both types of stands, nitrate reductase (NR) activity was detectable only in the period of vegetative growth and it was higher in the fragmented than in the closed stands. Glutamine synthetase (GS) activity showed three-fold higher activities in the leaves from the fragmented stands compared to those in closed stands, indicating high substrate (NH₃/NH₄⁺) availability. Polyamine concentrations were 4–10-fold higher in the leaves of the fragmented stands than in those of closed stands. Photosynthetic activity was nearly equal in both stands, however, photorespiration was about two-fold higher in the fragmented than in the closed stands. A linear correlation between photorespiration and GS activity indicated a causal relationship (R²=0.86). Stomatal conductance data suggest that the higher photorespiration in the fragmented stands could be the consequence of disturbed stomatal regulation. It is concluded that fragmented stands of Phragmites possess an altered C/N metabolism, due to high photorespiration and intensive N metabolism. The primary reason of the cascade of events is still not clear but apparently, these metabolic malfunctions accompany an accelerated die-back of Phragmites around Lake Balaton.     

Low levels of reserve carbohydrates in reed (Phragmites australis) stands of Kis-Balaton,Hungary

Seasonal dynamics of concentrations of reserve carbohydrates (starch, sucrose, glucose, fructose and their sum denoted as total non-structural carbohydrates, TNC) were followed in five reed stands of Kis-Balaton wetland area in Hungary. The stands included three stands of tall and robust reed, situated in the Ingói area, and two stands of short and subtle reed situated in the downstream part. While both the seasonal pattern and the proportions of single carbohydrate species corresponded to findings for other sites, the absolute concentrations were markedly lower as compared to other European reed stands except lake Fertö in Hungary. The seasonal minimum values ranged from 54 to 87 mg g⁻¹ dry weight. During the seasonal minimum, glucose reached the lowest concentrations (down to 0.01 mg g⁻¹ dry weight) of all carbohydrates measured. The seasonal maximum concentrations of TNC reached 160–270 mg g⁻¹ dry weight. Maximum TNC standing stock ranged from 240 to 520 g m⁻², and was depleted by 60–80% during the period of spring shoot growth. It is proposed that the remaining TNC pool may not suffice to support complete recovery after a subsequent catastrophic event.     

Detoxification of herbicides in Phragmites australis

Unintentional loss of herbicides into drainage ditches, shores or other waterbodies may cause large problems in farmland. Therefore strategies for the phytoremediation of agrochemicals and especially herbicides have become a topic of great interest in many agricultural areas. However, in order to establish effective biological pollution control, information on the detoxification capacity of riparian plants and aquatic macrophytes (e.g., Phragmites australis) is important to build up effective buffer stripes. We determined the detoxification capacity of Phragmites australis roots and leaves for the conjugation of agrochemicals to glutathione by assaying the model substrate CDNB as well as the herbicides fenoxaprop-P, propachlor, pethoxamid and terbuthylazine. Specific GST activities were always higher in the rhizomes (6.78 +/- 0.88 microkat/mg protein for CDNB) than in leaves (1.08 +/- 0.21 microkat/mg protein). The detoxification capacity is distributed across an array of GST isoforms. In summary, Phragmites australis seems to be efficient in herbicide detoxification and a good candidate for phytoremediation of effluents from agricultural sites.     

盐度变化条件下芦苇对互花米草的化感效应

以芦苇凋落物为试材,采用溶液浸提法和厌氧腐解法,获得水提物和腐解产物两种浸提物母液;在5‰和10‰盐浓度下以两种浸提物母液和25%母液分别处理互花米草种子、刚萌发的幼苗以及互花米草滩涂中特有的互花米草益生菌,考察盐度变化对互花米草种子的萌发、幼苗生长以及菌落生长的影响.结果表明：盐处理能够显著促进互花米草种子的萌发、幼根以及互花米草益生菌的生长(P＜0.05).较高的盐浓度(10‰)下,芦苇水提物对互花米草的萌发、生长和互花米草益生菌的生长呈促进作用;当盐浓度降低至5‰时,该促进作用消失.在较低的盐浓度(5‰)下,芦苇腐解产物对互花米草萌发产生显著的抑制作用;在较高盐浓度(10‰)时,则对互花米草益生菌产生显著促进作用(P＜0.05);在试验的任一盐浓度下,芦苇腐解产物对互花米草幼苗和幼根的生长均存在显著的抑制效果(P＜0.05).     

Between- and within-population differences in Phragmites australis

Phragmites australis (common reed) is a dominant clonal species in the interface between land and water in many European wetlands. Along the land-water gradient, strong consistently different selective forces might operate to give rise to genetic substructuring. I have investigated the occurrence of genetic substructuring in European P. australis populations. The present paper examines whether seedlings, from seeds collected at both ends of the land-water gradient, showed differences in response to nutrient supply. Under controlled conditions, the relative growth rate (RGR) in the exponential growth phase, and growth characters of 10-week old seedlings were assessed. Among populations, no differences in response to nutrient supply were found. Although total dry weight was not related to the geographic origin of the populations, northern/western compared to southern/eastern European populations (1) formed more but shorter shoots, (2) formed thinner but longer rhizomes, and (3) invested more dry matter in leaves at the expense of stems. It was concluded that these trait differences are likely to originate from differences in the length of the growing season in the native habitat. Within populations, ’water-side’ seedlings had a higher RGR under sub-optimal while for ’land-side’ seedlings this was under optimal nutrient conditions. Ten-week-old ’water-side’ seedlings had a higher total dry weight than ’land-side’ ones, irrespective of nutrient loading. Differences in growth could not clearly be related to differences in single biomass allocation and morphological traits. A discriminant analysis on these traits, however, revealed that ’water-side’ seedlings showed higher plasticity in discriminant scores than ’land-side’ seedlings in response to nutrient supply. Discriminant scores also pointed towards a subtle trade-off between height versus expansion growth of seedlings, from the water to landward side. In the Romanian population, this could be related to morphological differences between ploidy levels. Overall, it was concluded that within populations, selection on growth form rather than on adaptations to the nutrient status of the habitat might have taken place. Received: 20 August 1998 / Accepted: 29 July 1999     

莫莫格湿地芦苇生理生态特征对水深梯度的响应

通过在莫莫格湿地测量芦苇的生态特征、叶绿素含量和光化学效率,对水深梯度影响芦苇生理生态的规律进行了分析.芦苇株高、生物量、叶绿素含量以及最大光化学效率与湿地水深呈显著正相关关系,这些指标随水深的增加而提高;芦苇盖度与水深呈负相关关系,随着水深增加,芦苇盖度逐渐降低;芦苇最大光化学效率在6～7月份达到最大,进入8月后下降,同时芦苇生长减慢.结果表明,6～7月份是芦苇生长的关键时期,应尽量满足芦苇需水要求.15 cm水深是芦苇最小生态水位,而35～45 cm的水深能确保芦苇最佳生长状态,是芦苇生长的最适宜生态水位.     

Isoprene emission and primary metabolism in Phragmites australis grown under different phosphorus levels

Aquatic plants are generally used for wastewater purification and phytoremediation, but some of them also emit large amounts of isoprene, the most abundant biogenic volatile organic compound. Since isoprenoid biosynthesis requires high amounts of phosphorylated intermediates, the emission may also be controlled by inorganic phosphorus concentration (Pi) in leaves. We carried out experiments to determine the emission of isoprene from Phragmites australis plants used in reconstructed wetlands to phytoremediate elevated levels of phosphorus contributed by urban wastes. Four groups of plants were grown hydroponically in water containing different levels of KH(2)PO(4). High levels of phosphorus in the water resulted in high Pi in the leaves. High Pi stimulated photosynthesis at intercellular CO(2) concentrations lower and higher than ambient, implying higher ribulose 1,5-bisphosphate carboxylase (Rubisco) activity and higher ribulose 1,5-bisphosphate regeneration rates, respectively. However, isoprene emission was substantially lower at high Pi than at low Pi, and was not associated to photosynthesis rates at high Pi. This surprising result suggests that isoprene is limited by processes other than photosynthetic intermediate availability or by energetic (ATP) requirements under high Pi levels. Irrespective of the mechanism responsible for the observed reduction of isoprene emission, our results show that Phragmites plants may effectively remove phosphorus from water without concurrently increase isoprene emission, at least on a leaf area basis. Thus, Phragmites used in reconstructed wetlands for phytoremediation of urban wastes rich of phosphates will not contribute high loads of hydrocarbons which may influence air quality over urban and peri-urban areas.     

Phenotypic and genotypic variation of Phragmites australis: Comparison of populations in two human-made lakes of different age and history

Populations of Phragmites australis (CAV.) TRIN. ex STEUD. were studied in the littoral zones of two man-made lakes located in the Třeboň Basin (South Bohemia, the Czech Republic): (1) Opatovický fishpond, a shallow artificial lake constructed in 1510–1514 by damming a shallow valley and used since for carp production, and (2) Halámky sand pit, a new lake formed by sand extraction in 1970–1994. Phenotypic variability was assessed on the basis of shoot morphological and growth characteristics, measured at the time of seasonal maximum aboveground biomass. Genotypic variability was detected using RAPDs, which demonstrated a high clonal diversity in both habitats. The clonal diversity would be strongly underestimated if it were based only on morphological differences. Higher genotypic variability was found in the fishpond reed, not corresponding with low variability in its phenotype performance. Based on analysis of 160 samples, four patterns of genotypic variation were detected: (1) Some stands were genetically uniform and were therefore considered to be monoclonal in both populations studied. (2) Some stands consisted of several clones at the Halámky sand pit. However, these clones showed more similarity within the particular stands than with clones of adjacent stands. (3) In the Opatovický fishpond population, multiclonal stands consisted of clones with a low degree of similarity. (4) Identical clones were detected in several neighbouring stands separated by gaps in the Opatovický fishpond population. The findings support a model of colonization postulating that populations initiated by seeds are initially genetically diverse and over time become dominated by a few clones as a result of competition and selection. These processes then decrease both genetic and morphological variability.     

茭草和芦苇叶片光合与荧光特性对土壤持续干旱的响应

对茭草Zizania latifolia和芦苇Phragmites australi进行自然持续干旱处理,测定植物叶片光合作用特征参数和叶绿素荧光参数及相应的土壤水分含量,探讨茭草、芦苇叶片对土壤持续干旱胁迫的光合生理响应。随着土壤水分逐渐减少,茭草叶片净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)均先增大后减小,芦苇叶片的Pn、Tr、Gs均逐渐减小;茭草和芦苇叶片的胞间二氧化碳浓度(Ci)、光化学最大量子效率(Fv/Fm)、非光化学猝灭系数(NPQ)、水分利用率(WUE)、有效荧光产量(Yield)、电子传递速率(ETR)、光化学猝灭系数(qP)变化趋势一致,其中Ci逐渐增大,Fv/Fm、NPQ、WUE先增大后减小,Yield、ETR、qP逐渐减小。分析表明,当茭草和芦苇遭遇较严重干旱时,叶片Pn下降主要是非气孔限制因素导致;当生长基质(沙土)的田间持水量为19%,速效氮、磷、钾分别为28.14、2.25、25.76 mg·kg-1,pH 7.2时,茭草的干旱胁迫阈值在土壤含水量(2.17 ± 0.18)%和(1.66 ± 0.23)%之间,芦苇的干旱胁迫阈值在土壤含水量(1.84 ± 0.17)%和(0.25 ± 0.07)%之间。     

芦苇生态型研究进展

芦苇生态幅极广,适生于多种生境类型。不同的环境选择压力如水深、盐度、养分、气候等交互影响致使芦苇个体及种群间发生不同程度的分化和变异,形成了形态、生理或遗传上互有差异、异地性的不同生态型。尽管基于芦苇表型变异以及遗传变异进行生态型划分的研究已开展很多,但针对芦苇生态型变异规律及其可能的形成机制的认知仍存在较多分歧。在总结近年来有关芦苇生态型研究文献的基础上,通过对影响芦苇生态型变异的主要因素——环境因素和遗传因素的分析,以期为芦苇生态型的划分及其可能的形成机制提供新的研究思路。(1)空间尺度的选择应成为研究者分析、划分芦苇生态型的首要定位。在较大的地理空间尺度上,高度异质性的生境导致某些性状的变异式样具有相对的不连续性,可作为不同芦苇生态型鉴别与描述的主要依据;(2)在合理的尺度定位、取样设计和统计分析的基础上开展的表型变异研究,及进一步基于种群水平的分子标记研究(分子指纹特征或特有等位基因),可为芦苇生态型的鉴定、划分提供更为可靠的参考数据,并且可以甄别生境差异(环境响应)和遗传变异对芦苇不同生态型分化的贡献;(3)应同时进行不同生态型的特定性状与功能(株高、茎粗、生物量、生理抗逆性、水体污染物净化能力等)的定位,推动优良基因型的选育与扩繁。     

Phragmites australis and silica cycling in tidal wetlands

Tidal marshes have recently been shown to be important biogenic Si recycling surfaces at the land–sea interface. The role of vegetation in this recycling process has not yet been quantified. In situ and ex situ decomposition experiments were conducted with Phragmites australis stems. In a freshwater tidal marsh, litterbags were incubated at different elevations and during both winter and summer. Biogenic Si (BSi) dissolution followed a double exponential decay model in the litterbags (from ca. 60 to 15 mg g⁻¹ after 133 days), irrespective of season. Si was removed much faster from the incubated plant material compared to N and C, resulting in steadily decreasing Si/N and Si/C ratios. Ex situ, decomposition experiments were conducted in estuarine water, treated with a broad-spectrum antibiotic, and compared to results from untreated incubations. The bacterial influence on the dissolution of dissolved Si (DSi) from P. australis stems was negligible. Although the rate constant for dissolved Si dissolution decreased from 0.004 to 0.003 h⁻¹, the eventual amount of BSi dissolved and saturation concentration in the incubation environment were similar in both treatments. P. australis contributes to and enhances dissolved Si recycling capacity of tidal marshes: in a reed-dominated small freshwater tidal marsh, more than 40% of DSi export was attributable to reed decomposition. As the relation between tidal marsh surface and secondary production in estuaries has been linked to marsh Si cycling capacity, this provides new insight in the ecological value of the common reed.     

Physiological responses of Phragmites australis to wastewater with different chemical oxygen demands

Constructed wetlands have been widely used to treat various wastewaters with large differences in their concentration of pollutants. The capability of wetland plants to resist these wastewaters is crucial for a wetland's healthy development. Phragmites australis has been shown to have the capability to grow in simulated wastewater containing a wide concentration of pollutants. In this study, the physiological responses of P. australis to simulated wastewaters with high chemical oxygen demands (CODs) were investigated in a bucket experiment. P. australis was incubated in buckets for 30 days at five treatments of 0, 100, 200, 400, and 800 mg L^?1 COD simulated wastewater. The net photosynthesis rate of the plants declined markedly with increasing COD levels. Proline and malondialdehyde (MDA) contents also increased dramatically. The plants further showed a unimodal pattern of superoxide dismutase (SOD) and peroxidase (POD) distribution along external COD values on the whole, indicating that high COD values (≥200 mg L^?1) can disrupt the normal metabolism of the plant. High COD levels (COD ≥ 400 mg L^?1) caused evident physiological changes in P. australis.     

Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

北京地区芦苇资源状态及其多样性

为探明北京地区芦苇(Phragmites australis)的资源状态和多样性, 实地考察北京主要河流、湿地和水库, 发现北京地区芦苇总生长面积已超过600 hm²。芦苇染色体倍性以八倍体为主, 四倍体次之。在面积较大的湿地内, 八倍体单一芦苇群落占据优势地位; 而在城市的浅河内有形态和遗传性多样的混合种群。研究表明, 植物性状和倍性水平之间无显著相关性。在小清河发现了6种形态各异的芦苇克隆, 均属于叶绿体DNA片段的P单倍型; 其单倍体基因组大小为(0.499±0.019) pg, 变异系数为3.8%。这表明表型与单倍型之间也不具相关性。此外, 发现1个具有变叶特性的芦苇, 将其命名为金条芦苇。北京地区芦苇形态和遗传多样性为研究芦苇基因型与环境适应性之间的关系提供了珍贵的资源。     

Phenotypic and genotypic variation of Phragmites australis: A comparative study of clones originating from two populations of different age

Populations of common reed (Phragmites australis [Cav.] Trin. Ex Steud.) were studied in the littoral zones of two human-made lakes in the Třeboň Basin (South Bohemia, Czech Republic): (1) Opatovický fishpond, a 500-year-old fishpond, and (2) Halámky sand pit, a new lake formed by sand extraction in the 1970–1990s. Clones were identified using the RAPD method within morphologically different stands. The clones were propagated vegetatively and cultivated under comparable conditions in a common garden experiment for one vegetation season. Morphological and growth characteristics of the cultivated plants were recorded and compared with those found in the original reed stands. The younger (30-year old) population was genotypically more diverse than the older (500-year old) population. In addition, the younger population demonstrated correspondence of genotypic and phenotypic variation, while the older did not. The results indicate that the genetically determined morphological features (especially shoot length) are manifested more fully in the phenotype of the plants of the younger (Halámky) population, whereas environmental factors affect the phenotype to a greater extent in the older (Opatovický) population.     

丛枝真菌对互花米草和芦苇氮磷吸收的影响

互花米草(Spartina alterniflora Loisel.)是我国海滨盐沼的入侵植物,与土著种芦苇(Phragmites australis)形成了广泛的竞争;已知丛枝菌根(AMF)对不同植物的生长存在差异性影响;但其对互花米草与芦苇之间的种间关系,是否对互花米草入侵芦苇群体产生作用值得探讨.研究对两物种进行了丛枝菌根接种处理,种植模式处理和盐度处理的三因素实验.结果表明:盐度增加使得单种时芦苇、混种时互花米草的AMF侵染率显著下降(p<0.05),而混种时芦苇和单种时的互花米草AMF侵染率受盐度影响不显著(p>0.05).混种时,两种植物的丛枝菌根形成均受对方影响,并且盐度升高使两种植物之间对AMF侵染率的影响发生变化,在淡水生境下混种时,芦苇的AMF侵染率比单种时降低40.5%,互花米草的AMF侵染率比单种时提高了86.9%,均差异显著(p<0.05);在低盐度下混种时芦苇的AMF侵染率比单种时降低24.7%,差异显著(p<0.05),而对互花米草的影响不显著;在高盐度下混种对芦苇的AMF侵染率影响不显著,而使互花米草的AMF侵染率显著降低,降低率比例达78.7%.在淡水生境下,丛枝菌根对芦苇和互花米草的N、P吸收均有显著的促进作用;但是在咸水生境下生长时芦苇的N、P含量主要受盐度的显著影响(p<0.05),随盐度增加而增加;虽然在咸水生境下丛枝菌根仍旧促进芦苇的N、P吸收,但其影响远小于盐度的影响,并且促进效果受到盐度的抑制;但互花米草的N、P含量不受盐度影响.由此可见,接种AMF对这两种植物的氮磷吸收有着不同程度的促进,其作用大小与侵染程度有关,且受到盐度和种植模式的影响.