Nutrient concentrations in a Littorella uniflora community at higher CO2 concentrations and reduced light intensities

1. Oligotrophic softwater lakes represent a special type of aquatic ecosystem with unique plant communities where generalisations from other aquatic plant communities to rising CO₂ in the water column may not apply. 2. In the present study, we set up large in situ mesocosms and supporting laboratory experiments with isoetid vegetation (Littorella uniflora) where water column CO₂ and light could be manipulated in order to test whether (i) light and CO₂ availability affect nutrient concentrations in isoetid vegetation, and (ii) if changes in light and CO₂ climate affect fluxes of inorganic nitrogen (N) and phosphorus (P) from sediment to water column, which potentially could result in increased growth of epiphytic algae. 3. The results showed that the standing stocks of phosphorus and nitrogen in the L. uniflora vegetation were significantly influenced by CO₂ concentration and light intensity. Both standing stocks of P and N were significantly higher in the mesocosm treatments with high CO₂ concentration than in those at low CO₂ concentration. Similarly, standing stocks of P and N enhanced with increasing light intensity. 4. Measurements of nutrient fluxes both in the field and the laboratory did not show any significant release of nutrients to the water column from plants or sediments at any of the light or CO₂ treatments. However, mats of epiphytic algae developed from the beginning of June to late September and caused a light reduction for the isoetid vegetation. 5. Increasing CO₂ concentrations in the water column may over time potentially result in a change in soft water plant communities.     

A multidisciplinary approach to understanding the recent and historical occurrence of the freshwater plant, Littorella uniflora

1. The vulnerability of softwater, oligotrophic lakes to eutrophication has caused the disappearance of many, if not most, of the unique isoetid plant communities. We tested whether the presence or disappearance of the isoetid Littorella uniflora (L.) could be predicted from environmental parameters, soil types and land use in the catchment area, and atmospheric nitrogen deposition. 2. We found that the topographic catchment area of a lake was an irrelevant unit to study effects of soil type and land use. Instead, using a GIS‐generated buffer zone around the lakes it proved feasible to classify 472 lakes into historical (if L. uniflora had disappeared) or recent (if L. uniflora was still present) Littorella lakes, based on soil type and land use. Our analysis showed that aeolian sand deposits and heath in the buffer zone favoured the presence of L. uniflora, whereas moraine clay and agriculture were strongly linked to the disappearance of L. uniflora. 3. However, in order to understand fully the presence or disappearance of L. uniflora, environmental data were needed in addition to soil types, land use and nitrogen deposition, and the use of discriminant analysis allowed us to classify 96% of the investigated lakes correctly into recent or historical sites. Alkalinity, total phosphorus, total nitrogen, aeolian sand deposits and heath were the most important parameters explaining the presence or disappearance of L. uniflora. Our analysis also indicated that eutrophication, rather than acidification, has likely caused the disappearance of L. uniflora from 218 of the 472 lakes investigated. 4. Our findings have widespread implications for the conservation or restoration of isoetid habitats and we recommend applying a wide buffer zone around lakes, with restrictions on farming and traditional forestry activities. In addition, our buffering concept may prove a useful tool for aquatic ecologists to investigate relationships between catchment features and organisms (plants, insects and amphibians) with aquatic as well as terrestrial life forms.     

Growth rates and morphological adaptations of aquatic and terrestrial forms of amphibious Littorella uniflora (L.) Aschers.

Morphological – anatomical features of the terrestrial and the aquatic life form of the rosette species Littorella uniflora, inhabiting nutrient poor soils of oligotrophic lakes, were investigated together with growth rates of both life forms and of transplants. Growth rates were the same for the two life forms. However, growth of transplanted plants was somewhat reduced by transition from one environment to another. This was especially true for aquatic plants, which may be stressed by desiccation when moved to the terrestrial environment. The morphological – anatomical differences between the life forms were small compared with many other amphibious species which produce highly specialized leaves and life forms in air and under water. It is suggested that the conservative leaf morphology of Littorella is a consequence of the high dependence on rhizospheric CO₂ of both the aquatic and the terrestrial form of Littorella, making production of leaves specialized for carbon uptake in one specific environment unnecessary.     

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.     

Land plants of amphibious Littorella uniflora (L.) Aschers. maintain utilization of CO2 from the sediment

Summary Submerged macrophytes of the isoetid life form derive the majority of their CO₂ for photosynthesis from the sediment. The experiments described here were designed to test the hypothesis that root uptake of CO₂ is important also in the terrestrial form of Littorella uniflora. The results of ¹⁴CO₂ experiments showed that sediment CO₂ contributed 56% of the total fixation at 0.1mm CO₂ in the rhizosphere, 83% at 0.5mm and 96% at 2.5mm. Sediment CO₂ in emergent Littorella stands ranged from 0.1 to 1.0mm and averaged 0.5mm. Measurements of the net CO₂ exchange over the leaves showed an even higher dependence of the sediment as CO₂ source. Littorella leaves had no stomata at the base and densities (ca. 100 mm^–2) typical of terrestrial plants at the tip, allowing sediment-derived CO₂ to be supplied along the length of the leaf. The stomata permit supply of CO₂ from the air during periods of reduced sediment CO₂ concentrations (e.g. if the sediment dries up) and regulate transpiration.     

Massive uprooting of Littorella uniflora (L.) Asch. during a storm event and its relation to sediment and plant characteristics

During spring storms massive uprooting of Littorella uniflora occurred in a shallow Dutch softwater lake. The aim of this study was to test whether changes in plant morphology and sediment characteristics could explain the observed phenomenon. Uprooting was expected to occur in plants having a high shoot biomass and low root to shoot ratio (R:S), growing on sediments with a high organic matter content. Normally, uprooting of the relative buoyant L. uniflora is prevented by an extensive root system, expressed as a high R:S. This was studied by sampling floating and still rooted L. uniflora plants, as well as sediment and sediment pore water, along a gradient of increasing sediment organic matter content. Increasing organic matter content was related to increasing L. uniflora shoot biomass and consequently decreasing R:S. Furthermore, the results indicated that uprooting indeed occurred in plants growing on very organic sediments and was related to a low R:S. The increased shoot biomass on more organic sediments could be related to increased sediment pore water total inorganic carbon (TIC; mainly CO₂) availability. Additionally, increased phosphorus availability could also have played a role. The disappearance of L. uniflora might lead to higher nutrient availability in the sediments. It is suggested that this could eventually promote the expansion of faster‐growing macrophytes.     

独叶草叶宏观形态性状的变异式样的初步研究

根据独叶草分布区内 8个居群的叶的 8个宏观性状在散点图上的表现 ,探讨了独叶草的形态变异式样。结果表明 ,独叶草在叶宏观性状上表现出了一定的变异性 ,这种变异性表现在不同居群间及由不同居群组成的区域上 ,但在居群间或由不同居群组成的区域间在性状上有不同程度的重叠。     

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.     

Effects of arbuscular mycorrhizae on biomass and nutrients in the aquatic plant Littorella uniflora

1. It has been hypothesised that the symbiosis with arbuscular mycorrhizal fungi (AMF) leads to a higher uptake of phosphorus (P) and nitrogen (N) in aquatic plants, but it has never been shown experimentally without the use of fungicides. In particular, the symbiosis may be important for nutrient uptake by isoetids in oligotrophic lakes, where low concentrations of inorganic N and P both in the water and in the sediment limit the growth of plants and where symbiosis facilitates the uptake of nutrients from the sediment. 2. Plants of the isoetid Littorella uniflora were propagated under the sterile conditions without an AMF infection. The plants were then grown for 60 days with and without re‐infection by AMF, and with either high (150 μm ) or low (ambient concentration approximately 15 μm ) CO₂ concentration. 3. The study proved that the symbiosis between AMF and L. uniflora had a positive impact on the retention of N and P in the plants at very low nutrient concentrations in the water and on biomass development. Shoot biomass and standing stocks of both P and N were significantly higher in re‐infected plants. 4. Raised CO₂ concentration resulted in a fivefold increase in hyphal infection, but had no impact on the number of arbuscules and vesicles in the cross sections. There were significantly higher biomass and lower tissue P and N concentrations in the plants from high CO₂ treatments. This resulted in similar standing stocks of P and N in plants from low and high CO₂ treatments. 5. The results from this study showed that the symbiosis between AMF and L. uniflora is an important adaptation enabling isoetids to grow on nutrient‐poor sediments in oligotrophic lakes.     

苹果叶片气体交换日变化动态模拟

根据苹果叶片气体交换模型可模拟不同小气候条件下光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和水分利用效率(WUE)的日变化。Pn和Gs在晴天呈现出不对称双峰曲线:上午增加,约8:00前后达到最大值,然后下降,到16:00左右时再升高到第2个峰值,最后再下降。Tr晴天的日变化与Pn和Gs相似,在晴天呈"n"字形曲线,但是Tr的午休现象较Pn和Gs轻,并且最大值出现在午后。晴天时WUE在太阳出来不久就达到了最大值,并迅速下降到一个比较稳定的水平,傍晚前后降到0。模型模拟表明气体交换的日变化主要是由太阳辐射驱动的,而午休现象主要是由叶水势和湿度的减少引起。根据在富士苹果园观测数据(Maluspumila Mill.Fuji;北纬40°13′,东经116°13′,海拔79m)对模型进行了验证。结果表明在不同的有效光合辐射、温度、湿度和CO2浓度下Pn的观测值和模拟值非常吻合,树冠上层叶片Pn和Gs日变化的实测值和模拟值也基本一致。Tr的模拟值略有高估,可能是由叶片的相互遮荫及Pn和有效光合辐射之间的非线性关系引起。     

Reduced contribution from Na+/H+ exchange to acid extrusion during anoxia in adult rat hippocampal CA1 neurons

The effect of anoxia on Na+/H+ exchange activity was examined in acutely isolated adult rat hippocampal CA1 neurons loaded with the H+-sensitive fluorophore, BCECF. Five-minute anoxia imposed under nominally HCO3-/CO2-free conditions induced a fall in pHi, the magnitude of which was smaller following prolonged exposure to medium in which N-methyl-D-glucamine (NMDG+) was employed as an extracellular Na+ (Na(+)(o)) substitute. Also consistent with the possibility that Na+/H+ exchange becomes inhibited soon after the induction of anoxia, rates of Na(+)(o)-dependent pHi recovery from internal acid loads imposed during anoxia were slowed, compared to rates of Na(+)(o)-dependent pHi recovery observed prior to anoxia. At the time at which rates of pHi recovery were reduced during anoxia, cellular adenosine triphosphate (ATP) levels had fallen to 35% of preanoxic levels, suggesting that ATP depletion might contribute to the observed inhibition of Na+/H+ exchange. In support, incubation of neurons with 2-deoxyglucose and antimycin A under normoxic conditions induced a fall in cellular ATP levels that was also associated with reduced Na(+)(o)-dependent rates of pHi recovery from imposed acid loads; conversely, pre-treatment with 10 mm creatine attenuated the effects of anoxia to reduce both ATP levels and Na(+)(o)-dependent rates of pHi recovery from internal acid loads. Taken together, the results are consistent with the possibility that functional Na+/H+ exchange activity in adult rat CA1 neurons declines soon after the onset of anoxia, possibly as a result of anoxia-induced falls in intracellular ATP.     

蕤核叶片解剖结构和组织化学定位研究

采用解剖学和组织化学方法,研究了蕤核(Prinsepia uniflora Batal.)叶片的解剖结构,并对叶片中的黄酮类、生物碱类及多糖的组织化学定位进行了研究。结果表明:蕤核叶片上表皮有角质层,下表皮有气孔分布,气孔密度为278个·mm-2,近轴面栅栏组织细胞2～3层,排列紧密而整齐,含有许多晶体;叶片主脉木质部发达,由多列导管组成。上述特征说明蕤核叶片的解剖结构与环境之间的适应性。组织化学定位显示黄酮多分布于栅栏组织和厚角组织,生物碱含量少,多糖均匀分布于叶肉中。     

The behavioural responses and tolerance of freshwater benthic cyclopoid copepods to hypoxia and anoxia

The response of four benthic cyclopoid copepods,Acanthocyclops viridis (Megacyclops viridis) (Jurine, 1820),Macrocyclops albidus (Jurine),Eucyclops agilis (Koch, Sars) (Eucyclops serrulatus) (Fischer, 1851) andParacyclops fimbriatus (Fischer), to hypoxia and anoxia was investigated. All of these species died within six hours when confronted by totally anoxic conditions, but all survived four days at oxygen saturation levels as low as 25%. Males succumbed to the effects of anoxia more rapidly than the larger females of each species, and larger species survived for shorter periods than smaller species. In artificially stratified columns, where the lower layer was anoxic, all four species displayed an upward migratory response towards oxygenated conditions. Where the artificial hypolimnion was hypoxic, however, the migratory response was not observed. The results suggest that some benthic copepods cope with seasonal anoxia in eutrophic stratified lakes by migration rather than the various physiological adaptations shown by planktonic and semi-planktonic species.     

Contribution of the adaxial and abaxial surfaces of olive leaves to photosynthesis

Leaves of olive cultivars Frantoio and Maurino were irradiated with different irradiances from above, from below, or simultaneously from both directions to determine the contribution of the abaxial and adaxial leaf surfaces to photosynthesis. In both cultivars, irradiation of both sides of the leaf caused increases in net photosynthetic rate (P _N) and apparent quantum yield compared to irradiating only one surface with the equal photosynthetic photon flux density (PPFD), but the PPFD needed to saturate P _N decreased. At high and medium PPFD the P _N determined at irradiating both leaf surfaces was less than the sum obtained at irradiation of only the upper or the lower surface with the same PPFD. At PPFD higher than 1000 μmol m-2 s-1 in cv. Frantoio and 1200 μmol m-2 s-1 in cv. Maurino, P _N did not vary. At low PPFD (<200 μmol m-2 s-1), P _N at irradiating the adaxial and abaxial leaf surfaces simultaneously was about the sum of the values obtained by irradiating the upper and lower surfaces separately. Consequently the compensation irradiance was reduced from about 50 μmol m-2 s-1 to about 30 μmol m-2 s-1 when irradiating both leaf surfaces. The natural leaf orientation of the olive cultivar influenced the utilization of radiant energy by the abaxial surface. This revised version was published online in September 2006 with corrections to the Cover Date.     

Influence of quantity and lability of sediment organic matter on the biomass of two isoetids,Littorella uniflora and Echinodorus repens

1. Despite real improvement in the water quality of many previously eutrophic lakes, the recovery of submerged vegetation has been poor. This lack of recovery is possibly caused by the accumulation of organic matter on the top layer of the sediment, which is produced under eutrophic conditions. Hence, our objective was to study the combined effects of quantity and lability of sediment organic matter on the biomass of Echinodorus repens and Littorella uniflora and on the force required to uproot plants of L. uniflora. 2. Lake sediments, rich in organic matter, were collected from four lakes, two with healthy populations of isoetids and two from which isoetids had disappeared. The four lake sediments were mixed with sand to prepare a range of experimental sediments that differed in quantity and lability of sediment organic matter. Two isoetid species, E. repens and L. uniflora, were grown in these sediments for 8 weeks. Sediment quality parameters, including elemental composition, nutrient availability and mineralisation rates, were determined on the raw sources of sediment from the lakes. Porewater and surface water were analysed for the chemical composition in all mixtures. At the end of the experiment, plants were harvested and their biomass, tissue nutrient concentration and (for L. uniflora) uprooting force were measured. 3. For both species, all plants survived and showed no signs of stress on all types of sediment. The biomass of E. repens increased as the fraction of organic matter was increased (from 6 to 39% of organic content, depending upon sediment type). However, in some of the sediment types, a higher fraction of organic matter led to a decline in biomass. The biomass of L. uniflora was less responsive to organic content and was decreased significantly only when the least labile sediment source was used to create the gradient of organic matter. The increase in shoot biomass for both species was closely related to higher CO₂ concentrations in the porewater of the sediment. The force required to uproot L. uniflora plants over a range of sediment organic matter fitted a Gaussian model; it reached a maximum at around 15% organic matter and declined significantly above that. 4. Increasing organic matter content of the sediment increased the biomass of isoetid plants, as the positive effects of higher CO₂ production outweighed the negative effects of low oxygen concentration in more (labile) organic sediments. However, sediment organic matter can adversely affect isoetid survival by promoting the uprooting of plants.     

金钟藤叶片的气体交换特性

用LI-6400便携式光合测定系统(Li-CorInc.,USA)对广州林区新发现的入侵杂草金钟藤(Merremiaboisiana)叶片的气体交换进行了测定。结果表明(1)净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(E)的日变化均表现为单峰型曲线,中午金钟藤未出现光合作用抑制;最大光合速率可达20μmolCO2.m-2.s-1左右,Pn日平均值为8.8±0.75μmolCO2.m-2.s-1。(2)金钟藤光合作用的光饱和点较高,为1000~1200μmolphotons.m-2.s-1,表现出比较典型的阳生植物的特性。结果表明,金钟藤在自然环境中具有快速生长的特性可能与其较强和较稳定的光合能力有关。研究的初步结果对了解金钟藤迅速生长、具有高生产力和强大入侵力的原因提供了进一步深入探讨的思路和基础数据。     

Water‐level fluctuations affect sediment properties,carbon flux and growth of the isoetid Littorella uniflora in oligotrophic lakes

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