Adaptations for an amphibious life: changes in leaf morphology, growth rate, carbon and nitrogen investment, and reproduction during adjustment to emersion by the freshwater macrophyte Littorella uniflora

Littorella uniflora (L.) Ascherson is a small, perennial, amphibious rhizophyte of rosette life-form which is common along the margins of lakes, tarns and reservoirs where water-level fluctuations are often rapid and unpredictable. The majority of plants are continuously submersed and reproduce vegetatively, but a small proportion become completely emersed for variable lengths of time, when flowering and seed set occur. To find out how L. uniflora adjusts to sudden emersion we studied the plants at a reservoir where water level falls each spring and remains low throughout the summer; L. uniflora adjusted very quickly showing a degree of phenotypic plasticity not expected in a 'stress tolerator', including the production of a new set of terrestrial leaves with reduced lacunal volume and increased stomatal density, a rapid increase in leaf growth rate, and flowering within 3–4 wk. Comparison of terrestrial L. uniflora with aquatic plants growing permanently submersed in lake and tarn habitats showed that three to fourfold more carbon (C) and nitrogen (N) was incorporated into above-ground biomass by emersed plants. However, ramet production in the aquatic environment appeared to be more costly, in terms of C and N invested, than terrestrial flower and seed production. The combination of continuous, submersed vegetative spread with the capacity for a high degree of phenotypic plasticity allowing some flower and seed production to occur during brief periods of emersion seems to account for the success of this plant in the amphibious niche.     

BIOCHEMICAL HETEROPHYLLY AND FLAVONOID EVOLUTION IN NORTH AMERICAN POTAMOGETON (POTAMOGETONACEAE)

Morphologically heterophyllous species of Potamogeton also commonly display biochemical heterophylly with respect to flavonoid compounds. Generally, floating leaves contain an assortment of flavonoids, whereas submersed leaves often exhibit reduced flavonoid profiles. In strictly submersed (homophyllous) species, two patterns occur. Linear-leaved species have few flavonoids and their biochemical profiles resemble those of submersed leaves of heterophyllous species. Broad-leaved homophyllous species possess flavonoid profiles more similar to those of the floating leaves of heterophyllous species. Numerical analysis of these chemical data is consistent with phylogenetic relationships within the genus derived independently on the basis of morphological and chromosomal data. Glycoflavones, which are probably maintained in floating leaves because of their UV filtering ability, exhibit the most pronounced biochemical heterophylly in Potamogeton. The lack of glycoflavones in submersed leaves of heterophyllous species and in linear-leaved homophyllous species is attributable to the ability of naturally colored water to significantly absorb harmful UV radiation. These observations provide strong support for earlier hypotheses suggesting the importance of flavonoid evolution in the conquest of exposed terrestrial habitats by plants.     

武汉月湖水生植被重建的实践与启示

通过生物操纵、浮水植物和植物浮岛原位净化、岸边人工湿地净化和科学种植等措施,2－3年内,在长江中游地区、面积为0.66km2、水质为劣Ⅴ类的重富营养湖泊武汉月湖重建了以菹草和伊乐藻为优势种的沉水植物先锋群落,2005年春季的沉水植物盖度达到45％,提高了透明度。利用水位低、透明度相对高的冬季种植伊乐藻、菹草等适合冬春季生长的沉水植物是建立沉水植物先锋群落的一个有效途径。重污染水体过低的透明度是重建沉水植被的主要限制因子,在低水位或较高透明度时期,水生植物可以耐受相对较高的污染负荷,并且在有条件降低水位时,会大大提高重建沉水植被的成功率。重建并保持长期稳定的水生植被必须截污,并把营养盐浓度削减到一定范围内。水生植被重建初期是最脆弱的时期,水位、污染负荷等生态因子的较大波动很容易破坏植物群落,在这一时期要尽量保持关键因子的稳定,注意控制浮萍的扩张,使形成的生态系统逐步走向成熟。清除鱼类对提高水的透明度有重要作用,但必须伴随着水生植被的恢复或重建才能维持长久的良好效果。     

Abscisic acid regulation of heterophylly in Marsilea quadrifolia L.: effects of R-(-) and S-(+) isomers

The plant hormone abscisic acid (ABA) induces a developmental switch in the aquatic fern Marsilea quadrifolia, causing the formation of aerial type characteristics, including the elongation of petioles and roots, a change in leaf morphology, the expansion of leaf surface area, and the shortening of the internodes. A number of ABA-responsive heterophylly (ABRH) genes are induced early during the transition. Using optically pure isomers of ABA, it was found that both the natural S-(+)-ABA and the unnatural R-(-)-ABA are capable of inducing a heterophyllous switch and regulating ABRH gene expression. When dose responses are compared, the unnatural ABA gives stronger morphogenic effects than the natural ABA at the same concentration, it is effective at lower concentrations, and its optimal concentration is also lower compared with the natural ABA. Deuterium-labelled ABA enantiomers were used to trace the fate of applied ABA and to distinguish the applied compound and its metabolites from the endogenous counterparts. In tissues, the supplied (+)-ABA was metabolized principally to dihydrophaseic acid, while the supplied (-)-ABA was converted at a slower rate to 7'-hydroxy abscisic acid. Treatment with either enantiomer resulted in increased biosynthesis of ABA, as reflected in the accumulation of endogenous dihydrophaseic acid. Taken together, these results suggest two distinct mechanisms of action for (-)-ABA: either (-)-ABA is intrinsically active, or its activity is due to the stimulation of ABA biosynthesis.     

Heterophylly in Ranunculus flabellaris: The Effect of Abscisic Acid on Leaf Anatomy

Ranunculus flabellaris Raf., the yellow water crowfoot, exhibitsstriking heterophylly between submerged and terrestrial leaves.Leaves produced under water are highly divided with numerousnarrow lobes and deep sinuses, whereas terrestrial leaves havefew broad lobes and shallow sinuses. When plants are submergedin a 25 µM solution of ABA, the typical transition fromterrestrial to submerged leaves is completely suppressed and,instead, terrestrial-like leaves are produced. Image analysistechniques show that, in addition to this modification of leafmorphology, leaves produced under ABA treatment possess surfaceand internal features characteristic of terrestrial leaf anatomy.This study provides evidence that the environmental factorsthat influence the morphological and anatomical expression ofheterophylly may act through endogenous ABA. Ranunculus flabellaris, yellow water crowfoot, ABA, heterophylly, leaf anatomy     

Adaptive phenotypic plasticity: the case of heterophylly in aquatic plants

Phenotypic plasticity may play a key role in the adaptation of organisms to changing environmental conditions. A special case of plasticity is represented by heterophylly, the ability of semi-aquatic plants to produce different types of leaves below and above water. Submerged leaves are thin and lack both a cuticle and stomata, whereas aerial leaves are thicker, cutinized and bear stomata. The striking variability in the submerged, floating and aerial leaves of heterophyllous aquatics has historically been considered a paradigmatic example of adaptive phenotypic plasticity. An extensive body of developmental and physiological research reveals that heterophylly is quite often mediated by similar environmental cues across diverse taxa, which may imply a common underlying mechanism. Patterns of plasticity in response to environmental cues in the laboratory are consistent with the hypothesis of individual adaptation to heterogeneous environments, and the distribution of this trait among phylogenetically related aquatic angiosperms suggests either convergent or parallel evolution in their descent from terrestrial ancestors. Yet, critical evaluations of the ecological and evolutionary significance of this trait are scarce. In this essay, we discuss the patterns of plasticity revealed by experimental manipulative studies of heterophylly in the context of the general problem of adaptive phenotypic plasticity, and suggest avenues for future research that are needed in assessing the ecological and evolutionary significance of this trait.     

Effects of invasive species on plant communities: an example using submersed aquatic plants at the regional scale

Submersed aquatic plants have a key role in maintaining functioning aquatic ecosystems through their effects on the hydrological regime, sedimentation, nutrient cycling and habitat of associated fauna. Modifications of aquatic plant communities, for example through the introduction of invasive species, can alter these functions. In the Sacramento-San Joaquin River Delta, California, a major invasive submersed plant, Brazilian waterweed Egeria densa, has become widespread and greatly affected the functionality of the submersed aquatic plant community. Rapid assessments of the distribution and abundance of this species are therefore crucial to direct management actions early in the season. Given the E. densa bimodal growth pattern (late spring and fall growth peaks), summer assessments of this species may indicate which and where other submersed species may occur and fall assessments may indicate where this and other species may occur in the following spring, primarily because the Delta’s winter water temperatures are usually insufficient to kill submersed aquatic plant species. We assessed community composition and distribution in the fall of 2007 and summer of 2008 using geostatistical analysis; and measured summer biomass, temperature, pH, salinity, and turbidity. In the fall of 2007, submersed aquatic plants covered a much higher proportion of the waterways (60.7%) than in the summer of 2008 (37.4%), with a significant overlap between the seasonal distribution of native and non-native species. Most patches were monospecific, and multispecies patches had significantly higher dominance by E. densa, co-occurring especially with Ceratophyllum demersum. As species richness of non-natives increased there was a significant decrease in richness of natives, and of native biomass. Sustained E. densa summer biomass negatively affected the likelihood of presence of Myriophyllum spicatum, Potamogeton crispus, and Elodea canadensis but not their biomass within patches. Depth, temperature and salinity were associated with biomass; however, the direction of the effect was species specific. Our results suggest that despite native and invasive non-native submersed plant species sharing available niches in the Delta, E. densa affects aquatic plant community structure and composition by facilitating persistence of some species and reducing the likelihood of establishment of other species. Successful management of this species may therefore facilitate shifts in existing non-native or native plant species.     

激素对水生植物生理生态的影响及其应用

激素代谢是植物传导信号和调节生长发育的重要途径.陆地植物五大类激素在水生植物中也有分布,尽管近年来环境污染导致水生植物衰退的问题日益得到重视,但水生植物激素的研究和应用却远滞后于陆生植物.在总结了近年来激素类物质在水生植物中的研究成果,分别从激素的种类、激素的生理生态作用、激素生物合成的途径及作用的部位和机制、激素之间的相互作用.激素类物质在实验和实践上的应用等进行了全面阐述,指出了水生植物激素生理生态学研究的发展方向,从利用激素类物质诱导水生植物抗性的表达,提高抗逆性,恢复水生植被,以及研究和开发适于水生植物生产和管理的生长调节剂等方面,就水生植物激素的进一步研究和应用进行了探讨.     

长江中游浅水湖泊水生植物氮磷含量与水柱营养的关系

水生植物组织内氮和磷(N和P)含量受到水体营养状况和植物生长状况影响。对长江中游江汉湖群不同营养水平湖泊中大型水生植物的N和P含量3个季度的研究表明,在不同生活型水生植物中,沉水植物主要分布在中营养到中富营养湖泊中,在富营养湖泊均无分布,浮叶和挺水植物在不同营养类型湖泊的沿岸带均有分布。N和P含量以沉水植物最高,浮叶植物次之,挺水植物最低。水生植物的N和P含量都达到或超过生长所需最低N和P阈值,代表性浮叶植物和沉水植物的N和P含量随着湖泊营养水平提高呈现规律性变化。湖泊5种常见的水生植物N和P含量与水柱中不同种类N和P浓度具有季节性相关:菱(TrapabispinosaRoxb.)春夏季P含量都与TP(总磷)和TDP(总溶解磷)明显相关,春季N含量与NH4—N(氨氮)明显相关;春季黄丝草(PotamogetonmaackianusA.Benn.)的P含量与TP明显相关,夏季与TDP明显相关,春季和夏季黄丝草和穗花狐尾藻(MyriophyllumspicatumL.)的N含量与TN(总氮)和TDN(总溶解氮)显著正相关,秋季成负相关;夏季芦苇(PhragmitescommunisTrin.)P含量与TP和TDP显著相关;春季芦苇和香蒲(TyphaorientalisPresl.)N含量与NH4N和NO2N(亚硝态氮)显著相关。       

Physiological and photosynthetic plasticity in the amphibious, freshwater plant, Littorella uniflora, during the transition from aquatic to dry terrestrial environments

The physiological and photosynthetic responses of Littorella uniflora (L.) Ascherson, an amphibious macrophyte of isoetid life form, to rapid and prolonged emersion onto dry land, was studied at a reservoir. Water relations were little affected in the short term, but declining water potential and turgor pressure indicated water stress after flowering. High leaf lacunal CO₂ concentrations suggested continued CO₂ uptake from sediments. In contrast, a switch from Crassulacean acid metabolism (CAM) to C₃ photosynthesis was indicated by much lower levels of ΔH⁺ (down minus dusk titratable acidity) and phosphoenolpyruvate carboxylase (PEPC) activity in new terrestrial leaves, 7–8‐fold higher activity of ribulose bisphosphate carboxylase oxygenase (Rubisco), and increased chlorophyll and soluble protein contents. Accumulated nitrate and amino acid pools were depleted, whereas storage of carbohydrates as soluble sugars, fructan and starch increased. Plant carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) declined, perhaps reflecting changes in C fixation processes, N metabolism, and source C and N. In leaves of plants grown half‐emersed for an extended period, contrasting activities of PEPC and Rubisco were found in submersed and emersed portions. Overall, L. uniflora showed considerable phenotypic plasticity, yet seemed to remain poised for re‐submersion; these characteristics could be adaptive in the unpredictable water margin habitat.     

干出和沉水不同条件下羊栖菜光合作用碳源获得机制比较

经济海洋褐藻羊栖菜(Hizikia fusiforme(Harv.)Okamura)低潮时常常周期性地暴露于空气中.为了认识这种海藻在潮汐循环背景下的光合特征,对其在高潮沉水和低潮干出不同条件下的光合作用碳素获得机制进行了比较.沉水时,羊栖菜主要利用海水中HCO3-作为外源无机碳源驱动光合作用;而在干出条件下,其光合作用的主要碳源为空气中的CO2.在这两种不同环境条件下,光合作用与pH值的关系不同:沉水状态时,羊栖菜在高pH值(10.0)下光合活性很弱;而在干出条件下,羊栖菜在高pH值时仍有较高的光合活性.然而,光合作用无论是在沉水还是在干出条件下,对外源碳源的获得都表现出对胞外碳酸酐酶(CA)强烈的依赖性,并且其光合速率都受周围环境中无机碳源水平的限制.此外,在沉水和干出两种环境条件下,羊栖菜光合作用都表现出对氧气的敏感性.这表明,在羊栖菜中,依赖胞外CA的碳源获得机制不能使细胞内CO2浓度提高到阻碍其光呼吸的程度.增加空气中或海水中无机碳的浓度,能促进羊栖菜的光合作用,进而增加这种海藻的水产养殖产量.     

Trends of Superoxide Dismutase and Soluble Protein of Aquatic Plants in Lakes of Different Trophic Levels in the Middle and Lower Reaches of the Yangtze River,China

A limnological study was carried out to determine the responses of superoxide dismutase (SOD) activities and soluble protein (SP) contents of 11 common aquatic plants to eutrophication stress.Field investigation in 12 lakes in the middle and lower reaches of the Yangtze River was carried out from March to September 2004.Our results indicated that nonsubmersed (emergent and floating-leafed) plants and submersed plants showed different responses to eutrophication stress.Both SOD activities of the non-submersed and submersed plants were negatively correlated with their SP contents (P < 0.000 1).SP contents of non-submersed plants were significantly correlated with all nitrogen variables in the water (P < 0.05),whereas SP contents of submersed plants were only significantly correlated with carbon variables as well as ammonium and Secchi depth (SD) in water (P<0.05).Only SOD activities of submersed plants were decreased with decline of SD in water (P < 0.001).Our results indicate that the decline of SOD activities of submersed plants were mainly caused by light limitation,this showed a coincidence with the decline of macrophytes in eutrophic lakes,which might imply that the antioxidant system of the submersed plants were impaired under eutrophication stress.     

Osmotic stress, endogenous abscisic acid and the control of leaf morphology in Hippuris vulgaris L

Abstract. Previous reports indicate that heterophyllous aquatic plants can be induced to form aerial-type leaves on submerged shoots when they are grown in exogenous abscisic acid (ABA). This study reports on the relationship between osmotic stress (e.g. the situation encountered by a shoot tip when it grows above the water surface), endogenous ABA (as measured by gas chromatography-electron capture detector) and leaf morphology in the heterophyllous aquatic plant, Hippuris vulgaris. Free ABA could not be detected in submerged shoots of H. vulgaris but in aerial shoots ABA occurred at ca. 40ng (g fr wt)⁻¹. When submerged shoots were osmotically stressed ABA appeared at levels of 26 to 40ng (g fr wt)⁻¹. These and other data support two main conclusions: (1) Osmotically stressing a submerged shoot causes the appearance of delectable levels of ABA. (2) The rise of ABA in osmotically stressed submerged shoots in turn induces a change in leaf morphology from the submerged to the aerial form. This corroborates the hypothesis that, in the natural environment, ABA levels rise in response to the osmotic stress encountered when a submerged shoot grows up through the water/air interface and that the increased ABA leads to the production of aerial-type leaves.     

水体悬浮物浓度对大型沉水植物黑藻反射光谱的影响

沉水植物的恢复是改善富营养化水体和重建水生生态系统的有效措施．应用遥感技术可以实时、大面积监测沉水植物的分布和生长情况,而水体中的悬浮物等物质影响沉水植物的光谱特征,直接影响对湖泊、河流遥感影像中沉水植物的准确解译．本研究利用地物光谱仪在室外实测了不同水体悬浮物浓度条件下沉水植物黑藻群落的光谱反射率．结果表明,在400～900nm波段,水体中悬浮物浓度越高,沉水植物的光谱反射率越大．水体悬浮物浓度与黑藻群落光谱反射率在400～900nm均为显著相关（P〈0．05）,在450～900nm为极显著相关（P〈0．01）．在蓝、绿、红和近红外4个波段,将水体悬浮物浓度与其光谱反射率进行回归分析,均得到了较好的线性关系．利用本研究结果对富含悬浮物水体中沉水植物的反射光谱进行修正,可为准确、快速和大尺度遥感监测沉水植物的分布和动态提供科学依据．     

COMPONENTS OF PHOTOSYNTHESIS IN THE INTERTIDAL SACCATE ALGA HALOSACCION AMERICANUM (RHODOPHYTA,PALMARIALES)1

Rates of photosynthesis for the intertidal saccate alga Halosaccion americanum Lee were determined under submersed and emersed conditions. By fitting the data to a hyperbolic tangent function, P _max was 4.08 mmol CO₂. m^?2. h^?1 and I_k was 116.4 μE. m^?2. s^?1. under submersed conditions. Under emersed conditions, P _max was 1.89 mmol CO₂. m^?2. h^?1 and I_k was 22.9 μE. m^?2. s^?1. Dark fixation represented 3.7% of P_max in submersed thalli, whereas it equalled 33.3% of P_max in emersed thalli. Photosynthetic uptake from the thallus cavity represented a significant source of carbon, achieving 68.8% of that from the atmosphere and 29.4% of that from seawater. Retained seawater also greatly reduced drying under emersed conditions. Experimental thalli lost 70.4% of their water after 120 min under desiccating conditions, whereas control thalli lost only 6.3%. Emersed photosynthetic rates were enhanced by desiccation, At times, rates for desiccated thalli were two times those of fully-hydrated ones. Only after water loss exceeded 47% did photosynthetic rates fall below fully-hydrated rates. Utilizing data from this study a model was constructed to determine total photosynthetic production of H. americanum over a single daylight period. These caluclations demonstrate that photosynthetic contributions from emersed photosynthesis and retained seawater are significant. Because production from all sources is almost equal, total photosynthesis over a single day does not change greatly regardless of the time spent in air or in water.     

Correlations of Life Form,Pollination Mode and Sexual System in Aquatic Angiosperms

Aquatic plants are phylogenetically well dispersed across the angiosperms. Reproductive and other life-history traits of aquatic angiosperms are closely associated with specific growth forms. Hydrophilous pollination exhibits notable examples of convergent evolution in angiosperm reproductive structures, and hydrophiles exhibit great diversity in sexual system. In this study, we reconstructed ancestral characters of aquatic lineages based on the phylogeny of aquatic angiosperms. Our aim is to find the correlations of life form, pollination mode and sexual system in aquatic angiosperms. Hydrophily is the adaptive evolution of completely submersed angiosperms to aquatic habitats. Hydroautogamy and maleflower-ephydrophily are the transitional stages from anemophily and entomophily to hydrophily. True hydrophily occurs in 18 submersed angiosperm genera, which is associated with an unusually high incidence of unisexual flowers. All marine angiosperms are submersed, hydrophilous species. This study would help us understand the evolution of hydrophilous pollination and its correlations with life form and sexual system.     

Relative influences of submersed macrophytes and bioturbating fauna on biogeochemical processes and microbial activities in freshwater sediments

1. Invertebrates and aquatic plants often play a key role in biogeochemical processes occurring at the water–sediment interface of aquatic ecosystems. However, few studies have investigated the respective influences of plants and bioturbating animals on ecological processes (nutrient fluxes, benthic oxygen uptake, microbial activities) occurring in freshwater sediments. 2. We developed a laboratory experiment in aquaria to quantify the effects of (i) one invertebrate acting as a bioturbator (Tubifex tubifex); (ii) one submersed plant with a high sediment‐oxygenating potential (Myriophyllum spicatum) and (iii) one submersed plant with a low sediment‐oxygenating potential (Elodea canadensis). 3. The tubificid worms significantly increased the fluxes of nitrogen at the water–sediment interface (influx of nitrate, efflux of ammonium), whereas the two plant species did not have significant influences on these nitrogen fluxes. The differences in nitrogen fluxes between tubificid worms and plants were probably due to the bioirrigation process caused by T. tubifex, which increased water exchanges at the water–sediment interface. Tubifex tubifex and M. spicatum produced comparable reductions of nutrient concentrations in pore water and comparable stimulations of benthic oxygen uptake and microbial communities (percentages of active eubacteria and hydrolytic activity) whereas E. canadensis had a very weak influence on these variables. These differences between the two plants were due to their contrasting abilities to increase oxygen in sediments by radial oxygen losses (release of oxygen from roots). 4. Our study suggests that the bioirrigation process and radial oxygen loss are major functional traits affecting biogeochemical functioning at the water–sediment interface of wetlands.