Drag and reconfiguration of freshwater macrophytes

SUMMARY 1. Submerged freshwater macrophytes face large hydrodynamic forces in flowing waters in streams and on wave‐swept lake shores and require morphological adaptations to reduce the drag and the physical damage. This experiment studied five species of freshwater macrophytes and strap‐formed plastic leaves to test the predictions that: (i) increasing flexibility leads to greater reconfiguration and lower drag coefficients, (ii) flexible plants experience a steeper decline of drag coefficients with increasing water velocity than unflexible plants and (iii) plants mounted vertically on a horizontal substratum bend over in fast flow attaining a shielded position of low drag. 2. The results confirmed all three predictions. In fast flow, plants mounted upright on a horizontal platform gradually approached a position aligned with the flow, depending on their flexibility. In the range 8–50 cm s^?1 the deflection followed an interspecific negative linear relationship between log (tangent Φ) and velocity, where Φ represents the shoot angle normal to the horizontal level. Above 50 cm s^?1, further deflection was reduced perhaps by a combination of the elasticity and packing of shoots and the increasing lift generated by fast flow. 3. Drag coefficients of plants ranged between 0.01 and 0.1, typical of moderately to very streamlined objects. Drag coefficients declined log‐log linearly at increasing velocity, following negative slopes between ?0.67 and ?1.24 (median: ?1.0) because of reconfiguration and formation of a shielding canopy. Drag coefficients declined much less (median: ?0.55) for plants floating freely in the streaming water and which were capable of changing their shape but unable to form a shielding canopy. Drag coefficients declined even less for relatively unflexible plastic leaves (?0.30 to ?0.40), and they remained constant for stiff, bluff objects. 4. The experiments suggest that flow resistance of flexible, submerged macrophytes in natural streams may increase in direct proportion to water velocity because they form a shielding submerged canopy, and high water stages at peak flow may result in greater proportions of the water passing unimpeded above the canopy. In contrast, stiff amphibious and emergent reed plants should experience an increase of flow resistance with at least the square of velocity as reconfiguration is small and former aerial plant surfaces come into contact with the streaming water at higher water stages. Field experiments to test these predictions are urgently needed.     

An experimental study of habitat choice by Daphnia: plants signal danger more than refuge in subtropical lakes

1. In shallow temperate lakes, submerged plants often provide refuge for pelagic zooplankton against fish predation, a mechanism with potential strong cascading effects on water transparency and on the entire ecosystem. In (sub)tropical lakes, however, the interaction between aquatic plants and predation may be more complex, particularly because fish density is high within the plant beds in such systems. 2. Using laboratory ‘habitat choice’ experiments, we determined the effects of three (sub)tropical free‐floating plants, Eichhornia crassipes, Pistia stratiotes and Salvinia auriculata and the cosmopolitan submerged Ceratophyllum demersum, on horizontal movement by the water flea Daphnia obtusa. We tested for avoidance of plants in the absence and presence of alarm signals from crushed conspecifics and chemical cues from the fish Cnesterodon decemmaculatus, the fish have been subjected to different feeding regimes. 3. In the absence of other stimuli, D. obtusa strongly avoided the plants and the crushed conspecifics, as expected. However, the response to fish was insignificant regardless of their previous feeding regime. The avoidance of free‐floating plants was more pronounced than that of the submerged plant. Contrary to predictions based on research in temperate lakes, Daphnia did not take refuge among the plants but rather swam away from them when exposed simultaneously to plants and alarm signals. 4. We hypothesise that the avoidance of plants by D. obtusa may ultimately be attributable to an expectedly higher predation risk within the plants than in the pelagic, because of a high density of associated zooplanktivorous fish in the former. In the (sub)tropics, therefore, aquatic plants and particularly the free‐floating ones, may not promote cascading effects via Daphnia grazing on phytoplankton as seen in temperate eutrophic lakes.     

Transpiration does not control growth and nutrient supply in the amphibious plant Mentha aquatica

Mentha aquatica L. was grown at different nutrient availabilities in water and in air at 60% RH. The plants were kept at 600 mmol m^?3 free CO₂ dissolved in water (40 times air equilibrium) and at 30 mmol m^?3 CO₂ in air to ensure CO₂ saturation of growth in both environments. We quantified the transpiration-independent water transport from root to shoot in submerged plants relative to the transpiration stream in emergent plants and tested the importance of transpiration in sustaining nutrient flux and shoot growth. The acropetal water flow was substantial in submerged Mentha aquatica, reaching 14% of the transpiration stream in emergent plants. The transpiration-independent mass flow of water from the roots, measured by means of tritiated water, was diverted to leaves and adventitious shoots in active growth. The plants grew well and at the same rates in water and air, but nutrient fluxes to the shoot were greater in plants grown in air than in those that were submerged when they were rooted in fertile sediments. Restricted O₂ supply to the roots of submerged plants may account for the smaller nutrient concentrations, though these exceeded the levels required to saturate growth. In hydroponics, the root medium was aerated and circulated between submerged and emergent plants to minimize differences in medium chemistry, and here the two growth forms behaved similarly and could fully exploit nutrient enrichment. It is concluded that the lack of transpiration from leaf surfaces in a vapour-saturated atmosphere, or under water, is not likely to constrain the transfer of nutrients from root to shoot in herbaceous plants. Nutrient deficiency under these environmental conditions is more likely to derive from restricted development and function of the roots in waterlogged anoxic soils or from low porewater concentrations of nutrients.     

Plant distribution and abundance in relation to physical conditions and location within Danish stream systems

The distribution of obligate submerged plants, amphibious plants and terrestrial plants in streams was examined in relation to water depth, substrate and distance to the banks using univariate and mulitivariate analyses. The analyses were based on recordings in more than 40000 quadrats (25×25 cm) in 208 unshaded sites in the predominantly small and shallow (<1.6 m) Danish streams. Also, the distribution of plant abundance and richness from the source to the outlet of the stream system was determined.The submerged plants in Danish streams include 87 terrestrial, 22 amphibious and 30 obligate submerged taxa. The distribution of plant types was mainly related to water depth and distance to the bank among the physical conditions, while the type of bottom substrate had no significant influence. Terrestrial plants and amphibious plants (excluding Sparganium emersum) dominated in shallow water near the bank, but declined rapidly with increasing depth and distance to the bank, reflecting the importance of dispersal by ingrowth from populations on the banks to the water among these plants. Accordingly, these two plant groups constituted a higher proportion of total plant abundance in small streams than large streams. S. emersum dominated on great depth and distance to the banks, probably reflecting the lengthwise dispersal of this species from upstream to downstream parts of the stream system, the tolerance of the species to weed cutting and the adaptation to grow at low light intensities. Obligate submerged plants dominated at intermediate depths and at all distances from the bank except from 0 to 50 cm. This distribution reflects the ability of these species to disperse lengthwise in streams and live permanently submerged.The species number of all species and obligate submerged plants was lower in the smallest stream sites compared to larger downstream sites, while there was no difference for terrestrial and amphibious plants. The downstream increase of submerged species can be explained by the increase of habitat area and the dispersal of plants with the current, implying that the species pool accumulates with distance from the source. This result is at variance with a maximum richness of submerged plant species predicted for intermediate-sized streams according to the River Continuum Concept developed for large North American streams having forested shallow upstream sections and unshaded, deep downstream sections both unsuitable to submerged plant growth. The results for Danish streams imply that both the longitudinal connection through the flowing water and the transversal connection to the local species on the banks are important for plant distribution in the streams.     

水菜花异形叶片叶绿素荧光特征与HCO3-利用能力的研究

以生长发育过程中具有异形叶的水生植物水菜花（Ottelia cordata（Wall.）Dandy）为材料,利用叶绿素荧光技术和pH-漂移（pH-drift）实验测定具有两型叶片的水菜花正常沉水叶（submerged leaves,S）、正常浮水叶（floating leaves,F）、处于淹没状态的浮水叶（submerged floating leaves,SF）以及只有浮水叶植株的浮水叶（floating leaves only,OF）的叶绿素荧光特征与HCO₃^-利用能力。快速光曲线分析结果显示：沉水叶（S）的相对电子传递速率（rETR）显著低于其他类型的叶片,且随着光强的增强,差异增大;由光化学引起的荧光淬灭q_P、潜在最大电子传递速率（rETR_max）、半饱和光强（E_k）等指标均为沉水叶（S）显著低于浮水叶（F）。沉水叶（S）与淹没状态的浮水叶（SF）表征HCO₃^-利用能力的C_T/Alk值显著低于两种浮水叶（F和OF）。研究结果表明水菜花成年期的浮水叶比幼年期的沉水叶具有更高的光合效率,并且更适合在高光强下生长;当水位发生波动时被淹没的浮水叶会增强对HCO₃^-的利用能力以适应淹水低浓度CO₂环境。     

Morphological adaptations of emergent plants to water flow: a case study with Typha angustifolia, Zizania latifolia and Phragmites australis

1. Water velocity plays an important role in shaping plant community structure in flowing waters although few authors have yet attempted to explain the adaptation of plants to flow. 2. We aimed to test two hypotheses, that: (i) some emergent macrophytes reconfigure their shoot distribution in fast currents and form clumps, and (ii) the shape and morphology of such clumps minimises drag caused by the current. The study focuses on three emergent macrophytes that co‐occur along a gradient of water velocity. 3. The species showed a clear zonation in response to water depth and current velocity. Phragmites australis occupied shallower and more slowly flowing water than Typha angustifolia and Zizania latifolia, which had similar preferences. 4. Both T. angustifolia and Z. latifolia shoots were more clumped at high velocity, whereas they were more randomly distributed at low flow or in stagnant water. Because of the low shoot density, water flowed more easily through T. angustifolia clumps, whereas Z. latifolia clumps had a high shoot density and large amounts of trapped litter, causing stagnant water in the centre of the clump. The clumps of Z. latifolia with a high density of shoots were longer and narrower than T. angustifolia clumps. Phragmites australis was less tolerant of flow than the other two species and large amounts of litter trapped in the clumps impaired flow. 5. The shoot distribution of both T. angustifolia and Z. latifolia is reconfigured at high flow and this minimises drag on the clumps.     

Effects of spring acid episodes on macroinvertebrates revealed by population data and in situ toxicity tests

1. While streams in Europe and North America are now recovering chemically from chronic acidification, severe episodic acidification continues to threaten sensitive biota. To appraise further the biological importance of episodic acidification, we surveyed the distribution of the mayfly Baetis alpinus in streams in the Southern Alps (Canton Ticino, Switzerland) in relation to runoff acidity during spring floods. Moreover, to improve mechanistic understanding, in situ toxicity assays were carried out on nymphal B. alpinus during low flows and spring floods, both in streams prone to acid episodes and in well‐buffered controls. 2. Streams surveyed for invertebrates represented three groups which contrasted in susceptibility to episodic acidity. Group one included streams that were acid (alkalinity <0) in spring; group two streams were susceptible to acid episodes because of low base‐flow alkalinity (<200 μeq L^?1); and group three streams were well‐buffered and unlikely ever to be acid. The abundance of B. alpinus was similar among groups during stable flows in winter and summer, but was significantly lower in groups one and two following spring snowmelt. 3. During the bioassays, control streams remained circumneutral to alkaline (pH >6.4). By contrast, episodic streams were circumneutral at low flows, but became acid (pH 4.5–5.6 and total dissolved aluminium to 276 μg L^?1) during intense spring snowmelt. After 15‐day exposures, nymphal B. alpinus survival in the circumneutral control streams exceeded 92% irrespective of flow. In the episodic streams, survival matched the controls during low flows, but declined significantly to 10–20% during acid episodes in spring. Shorter exposure (2–4 days) to pH 5.5–5.7 did not cause significant mortality either during the exposure or over the following 7 days, indicating that B. alpinus might recover from short acid episodes. 4. Our data suggest that the spring distribution of B. alpinus in acid sensitive parts of the Alps directly reflects the toxicity of acid runoff during snowmelt. Our study illustrates that even mild episodic acidification can have significant consequences in Alpine streams for one of the most important invertebrate indicators of acidity.     

Temporal Persistence of Resource Polymorphism in Brook Charr, Salvelinus Fontinalis

Resource polymorphisms are believed to be an important intermediate step in competitive sympatric speciation. Reports of resource polymorphisms in fishes are becoming more common, but tests of the temporal persistence of resource polymorphisms remain scarce. We examined the temporal persistence of a body shape dimorphism reported previously for stream dwelling young-of-the-year brook charr, Salvelinus fontinalis, inhabiting slow vs. fast running water. In the year of our study, we found no difference in body shape between individuals in slow vs. fast flowing water, although fish in fast flowing water were of greater standard length than those in slower flowing water. We conclude that the resource polymorphism reported earlier for brook charr in streams is not stable temporally.     

Effects of habitat complexity on community structure and predator avoidance behaviour of littoral zooplankton in temperate versus subtropical shallow lakes

1. Structural complexity may stabilise predator–prey interactions and affect the outcome of trophic cascades by providing prey refuges. In deep lakes, vulnerable zooplankton move vertically to avoid fish predation. In contrast, submerged plants often provide a diel refuge against fish predation for large‐bodied zooplankton in shallow temperate lakes, with consequences for the whole ecosystem. 2. To test the extent to which macrophytes serve as refuges for zooplankton in temperate and subtropical lakes, we introduced artificial plant beds into the littoral area of five pairs of shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N). We used plants of different architecture (submerged and free‐floating) along a gradient of turbidity over which the lakes were paired. 3. We found remarkable differences in the structure (taxon‐richness at the genus level, composition and density) of the zooplankton communities in the littoral area between climate zones. Richer communities of larger‐bodied taxa (frequently including Daphnia spp.) occurred in the temperate lakes, whereas small‐bodied taxa characterised the subtropical lakes. More genera and a higher density of benthic/plant‐associated cladocerans also occurred in the temperate lakes. The density of all crustaceans, except calanoid copepods, was significantly higher in the temperate lakes (c. 5.5‐fold higher). 4. Fish and shrimps (genus Palaemonetes) seemed to exert a stronger predation pressure on zooplankton in the plant beds in the subtropical lakes, while the pelagic invertebrate Chaoborus sp. was slightly more abundant than in the temperate lakes. In contrast, plant‐associated predatory macroinvertebrates were eight times more abundant in the temperate than in the subtropical lakes. 5. The artificial submerged plants hosted significantly more cladocerans than the free‐floating plants, which were particularly avoided in the subtropical lakes. Patterns indicating diel horizontal migration were frequently observed for both overall zooplankton density and individual taxa in the temperate, but not the subtropical, lakes. In contrast, patterns of diel vertical migration prevailed for both the overall zooplankton and for most individual taxa in the subtropics, irrespective of water turbidity. 6. Higher fish predation probably shapes the general structure and dynamics of cladoceran communities in the subtropical lakes. Our results support the hypothesis that horizontal migration is less prevalent in the subtropics than in temperate lakes, and that no predator‐avoidance behaviour effectively counteracts predation pressure in the subtropics. Positive effects of aquatic plants on water transparency, via their acting as a refuge for zooplankton, may be generally weak or rare in warm lakes.     

Influence of submergence on growth of seedlings of Scirpus lacustris and Phragmites australis

• 1 Seeds of Scirpus lacustris and Phragmites australis were germinated in early June, and twenty-four seedlings of each species were subsequently exposed to submerged conditions (eight seedlings at each of the water depths 0.2, 0.4 and 0.8m), in outdoor 500–1 tanks in southern Sweden. Weight and shoot length of the plants were measured in September.
• 2 The Phragmites seedlings did not show any significant growth when submerged. The Scirpus seedlings, however, developed submerged leaves and exhibited considerable submerged growth. One Scirpus plant, in shallow water (0.2m), had developed an aerial shoot by September. Shoot length of the remaining (submerged) Scirpus plants was positively related to plant weight within water depth treatments, and was higher, in relation to plant weight, in deeper water. Mean weight in September of the submerged Scirpus plants decreased with increased water depth.
• 3 In south Swedish lakes with a lowered water table, Scirpus often occupies large areas on the lakeward side of the reed belt, which is generally dominated by Phragmites. The differences between the two species, in performance of submerged seedlings, suggest that this zonation may be created through successful submerged seedling establishment of Scirpus on the lakeward side of Phragmites.

     

Dynamic Properties of Recruitment,Expansion and Mortality of Macrophyte Patches in Streams

Streams are physically perturbed habitats with high demands on the dispersal and recruitment to maintain plant populations. Yet, little is known about these important demographic processes for stream plants. Therefore, we studied the monospecific vegetation of Callitriche cophocarpa in a small Danish lowland stream to determine: 1) the importance of drifting shoots and seeds for recruitment of plants, and 2) the influence of water flow, light availability and patch size on recruitment, growth and mortality processes. We found that the majority (about 90%) of new colonising patches of plant stands derived from drifting shoots being caught around protruding stones, while few developed from seeds. Many new patches were lost in the flowing water before roots became well established. Flow exposure of the patches resulted in the main growth taking place in the downstream direction. Combined areal cover of Callitriche patches on the stream bottom reached an upper limit of about 70%, probably because areal expansion above this threshold was constrained by strong shear forces and coarse substrata developing in the flow channels between the patches. We discuss why efficient shoot dispersal and vegetative growth documented here for Callitriche is an optimal plant strategy in flow-perturbed streams in contrast to the production of numerous small seeds but limited vegetative spread among ruderal plants in perturbed habitats on land.     

Strong growth limitation of a floating plant (Lemna gibba) by the submerged macrophyte (Elodea nuttallii) under laboratory conditions

1. The asymmetric competition for light and nutrients between floating and submerged aquatic plants is thought to be key in explaining why dominance by either of these groups can be stable and difficult to change. 2. Although the shading effect of floating plants on submerged plants has been well documented, the impact of submerged plants on floating plants has been poorly explored hitherto. 3. Here, we used laboratory experiments to examine how submerged plant (Elodea nuttallii) alter nutrient conditions in the water column and how this affects the growth of floating plants (Lemna gibba). 4. We demonstrate that, at higher nutrient concentrations, Lemna is increasingly likely to outcompete Elodea. 5. Under low nutrient concentrations (0.1–2 mg N L^?1) Elodea can strongly reduce the growth of Lemna. Growth of floating plants virtually stopped in some of the experiments with Elodea. 6. Extremely reduced tissue N, Mn, chlorophyll and elongated roots indicated that the growth inhibition of Lemna by Elodea was predominantly caused by the latter’s impact on the nutrient conditions for floating plants. 7. These results strengthen the hypothesis that submerged plants can prevent colonization of a lake by floating plants.     

Ethylene is required in tobacco to successfully compete with proximate neighbours

Plants sense neighbours even before these cause a decrease in photosynthetic light availability. Light reflected by proximate neighbours signals a plant to adjust growth and development, in order to avoid suppression by neighbour plants. These phenotypic changes are known as the shade‐avoidance syndrome and include enhanced shoot elongation and more upright‐positioned leaves. In the present study it was shown that these shade‐avoidance traits in tobacco (Nicotiana tabacum) are also induced by low concentrations of ethylene. Furthermore, it was shown that transgenic plants, insensitive to ethylene, have a delayed appearance of shade‐avoidance traits. The increase in both leaf angles and stem elongation in response to neighbours are delayed in ethylene‐insensitive plants. These data show that ethylene is an important component in the regulation of neighbour‐induced, shade‐avoidance responses. Consequently, ethylene‐insensitive plants lose competition with wild‐type neighbours, demonstrating that sensing of ethylene is required for a plant to successfully compete for light.     

Germination and early growth of Nymphaea odorata at different water depths

We experimentally determined the effects of water depth on seed germination and seedling growth and morphology, and we documented the transition from submerged to emergent plants in the white water lily, Nymphaea odorata. Seeds of N. odorata were germinated at 30, 60, and 90 cm water depth in outdoor mesocosms and percent germination and morphology measured after a month. The presence of self-seeded seedlings in pots at the same 3 water levels was also recorded over two years. To examine juvenile growth, seeds planted in soil were placed at the same mesocosm depths; germination and growth were monitored for three months, when the plants were harvested for morphological and biomass measurements. N. odorata germinated equally well in 30, 60 and 90 cm water; seedlings grew as submerged aquatics. After one month, seedlings in 90 cm water had less biomass than those in 30 cm (1.1 vs. 3.3 mg and 1.0 vs. 1.8 mg for different seed sources, respectively) and allocated relatively more biomass to shoots (97.5 vs. 67.8% and 73.1 vs. 58.0%, respectively). Seedlings in 60 cm water were intermediate. After 3 months of submerged growth, plant biomass remained less in 90 vs. 60 and 30 cm water (22.5 vs. 36.4 and 33.3 mg, respectively). Plants in 90 and 60 cm water had greater biomass allocation to shoots than plants in 30 cm water (85.7 and 72.6% vs. 64.4%, respectively) and produced larger laminae on longer petioles (lamina length = 33.3 vs. 25.2 mm in 90 vs. 30 cm; petiole length = 99.0 vs. 36.0 mm, respectively). After about 3 months, submerged plants produced floating leaves that had 39% shorter laminae but 267% to 1988% longer petioles than submerged leaves on the same plant. Lamina length to width allometric relations of submerged leaves were >1 at all water levels, distinguishing them from the equal allometry of adult floating leaves. The switch from production of submerged to emergent leaves resembles submergence-escape growth in other aquatics, but because the seedlings have been submerged throughout their life, submergence itself cannot be the stimulus to produce emergent leaves in these totally immersed plants. Our data show that N. odorata plants can establish from seeds in up to 90 cm water and that seedlings grow as submerged aquatics until they switch abruptly to production of floating leaves.     

The interaction of water flow and nutrients on aquatic plant growth

A long-term experiment was conducted to compare the effects of flowing and still water on growth, and the relationship between water flow and nutrients, in Aponogeton elongatus, a submerged aquatic macrophyte. A. elongatus plants were grown for 23 weeks with three levels of nutrition (0, 0.5 and 1g Osmocote Plus® fertiliser pot^–1) in aquaria containing stirred or unstirred water. Fertilized plants grew much better than non-fertilized. The highest fertilizer level produced 29% wider leaves and 58% higher total dry weight in stirred water. Stirred water increased leaf area by 40% and tuber size by 81%, but only with the highest level of nutrition. These results suggest that this plant depends on its roots for mineral uptake, rather than from the open water, and the major limitation to growth in still water is the supply of dissolved inorganic carbon. It was the combined effects of nutrient availability and stirring that produced the strongest response in plant growth, morphology and composition. This study provides some explanation for the observations of others that these plants grow best in creeks or river systems with permanently flowing water.     

Dispersal of plant fragments in small streams

1. Streams are subject to frequent natural and anthropogenic disturbances that cause sediment erosion and loss of submerged vegetation. This loss makes downstream transport and retention of vegetative propagules on the streambed very important for re‐establishing vegetation cover. We measured dispersal and retention of macrophyte stem fragments (15–20 cm long) along 300 m long reaches of four small to medium sized Danish lowland streams. 2. The number of drifting stem fragments declined exponentially with distance below the point of release. This finding makes the retention coefficient (k, m⁻¹) in the exponential equation a suitable measure for comparisons among different macrophyte species, and between stream reaches of different hydrology and vegetation cover. 3. Buoyancy of macrophyte tissue influenced retention. Elodea canadensis stems drifted below the water surface, and were more inclined to be retained in deeper water associated with submerged plants and obstacles in the streambed. Ranunculus peltatus stems were more buoyant, drifted at the water surface, and were more inclined to be trapped in shallow water and in riparian vegetation. 4. The retention coefficient of drifting stems increased with the relative contact between the flowing water and streambed, bank and vegetation. Thus, the retention coefficients were highest (0.02–0.12 m⁻¹) in shallow reaches with a narrow, vegetation‐free flow channel. Here there were no significant differences between E. canadensis and R. peltatus. Retention coefficients were lowest (0.0005–0.0135 m⁻¹) in deeper reaches with wider vegetation‐free flow channels. Retention of E. canadensis was up to 16 times more likely than retention of R. peltatus. 5. Overall, the longitudinal position in the stream system of source populations of species capable of producing numerous stems, the species‐specific retention coefficients of stems, and the retention capacity of stream reaches should be important for species distribution in perturbed stream systems. Retention of stems is probably constrained in headwaters by the small downstream flux of stem fragments because of the restricted source area, and constrained in downstream reaches by small retention coefficients. Macrophyte retention may, consequently, peak in medium‐sized streams.     

Effects of water velocity on photosynthesis and dark respiration in submerged stream macrophytes

The effects of flow velocities on dark respiration and net photosynthesis of eight submerged stream macrophytes were examined in a laboratory oxygen chamber. The shoots/leaves were exposed to saturating free-CO₂ concentrations and were attached basally so that they could move in the flowing water. Net photosynthesis declined by 34–61% as flow velocity increased from 1 to 8.6cm s^?1, while dark respiration increased 2.4-fold over the same range. The increase in dark respiration could only account for between 19 and 67% of the decrease in net photosynthesis. The relationship between flow velocity (U) and net photosynthesis (P) was described by: P=b×U^a. The exponent, a, varied from -0.20 to –0.48 and showed a negative correlation to the surface: volume (SA: V) ratio of the plants, i.e. species with high SA: V ratio were more sensitive to flow. In contrast, net photosynthesis of plants firmly attached to a supporting frame was not significantly affected by increasing flow velocity. This result indicates that the physical stress imposed on the plants by agitation or stretching in the flowing water is a key factor for the observed response.     

Can warm climate‐related structure of littoral predator assemblies weaken the clear water state in shallow lakes?

Shallow lakes, the most abundant lake type in the world, are very sensitive to climatic changes. The structure and functioning of shallow lakes are greatly impacted by submerged plants, and these may be affected by climate warming in various, contrasting, ways. Following a space‐for‐time substitution approach, we aimed to analyse the role of aquatic (submerged and free‐floating) plants in shallow lakes under warm climates. We introduced artificial submerged and free‐floating plant beds in five comparable lakes located in the temperate zone (Denmark, 55–57 °N) and in the subtropical zone (Uruguay, 30–35 °S), with the aim to study the structure and dynamics of the main associated communities. Regardless of differences in environmental variables, such as area, water transparency and nutrient status, we found consistent patterns in littoral community dynamics and structure (i.e. densities and composition of fish, zooplankton, macroinvertebrates, and periphyton) within, but substantial differences between, the two regions. Subtropical fish communities within the macrophyte beds exhibited higher diversity, higher density, smaller size, lower relative abundance of potentially piscivores, and a preference for submerged plants, compared with otherwise similar temperate lakes. By contrast, macroinvertebrates and cladocerans had higher taxon richness and densities, and periphyton higher biomass, in the temperate lakes. Several indicators suggest that the fish predation pressure was much stronger among the plants in the subtropical lakes. The antipredator behaviour of cladocerans also differed significantly between climate zones. Submerged and free‐floating plants exerted different effects on the spatial distribution of the main communities, the effects differing between the climate zones. In the temperate lakes, submerged plants promoted trophic interactions with potentially positive cascading effects on water transparency, in contrast to the free‐floating plants, and in strong contrast to the findings in the subtropical lakes. The higher impact of fish may result in higher sensitivity of warm lakes to external changes (e.g. increase in nutrient loading or water level changes). The current process of warming, particularly in temperate lakes, may entail an increased sensitivity to eutrophication, and a threat to the high diversity, clear water state.     

Sources of nutrients to rooted submerged macrophytes growing in a nutrient-rich stream

1. The relative contribution of roots and leaves to nutrient uptake by submerged stream macrophytes was tested in experiments where plants were grown in an outdoor flow-channel system. Water was supplied from a nutrient-rich stream with inorganic nitrogen and phosphorus concentrations typical of Danish streams.
2. Four submerged macrophyte species were tested, Elodea canadensis , Callitriche cophocarpa , Ranunculus aquatilis and Potamogeton crispus, and all species were able to satisfy their demand for mineral nutrients by leaf nutrient uptake alone. This was evident from manipulative experiments showing that removal of the roots had no negative impact on the relative growth rate of the plants. Further, the organic N and P concentrations of the plant tissue was constant with time for the de-rooted plants.
3. Enrichment of water and/or sediment had no effect on the relative growth rate of two species, E. canadensis and C. cophocarpa , indicating that in situ nutrient availability was sufficient to cover the needs for growth. Despite the lack of a response in growth rate, a reduced root/shoot biomass ratio was observed with nutrient enrichment of water and/or sediment, and an increased tissue-P concentration in response to open-water enrichment.
4. The open-water nutrient concentrations of the stream in which the experiments were performed are in the upper part of the range found for Danish farmland streams (the majority of Danish streams). Still, however, the negligible effect of nutrient enrichment on the growth of submerged macrophytes observed suggests that mineral nutrient availability might play a minor role in controlling macrophyte growth in most Danish streams.     

Effects of Air Contact on Growth,Inorganic Carbon Sources,and Nitrogen Uptake by an Amphibious Freshwater Macrophyte

Callitriche cophocarpa Sendtner is a heterophyllous amphibious macrophyte that produces apical rosettes of floating leaves. The importance of air contact for inorganic carbon and N uptake and for growth was investigated. Plants were grown with the floating rosette in contact with air of various humidities (10, 50, and >90% relative humidity) and with the submerged parts in N-free water at 350 [mu]M free CO2 and the roots in sediment with low or high NH3-N content. Humidity greatly affected the transpiration rate, whereas growth rate and N content were unaffected and were comparable to values measured for fully submerged shoots. Air contact had, however, a significant impact on growth when the free CO2 concentration in the water was low. Thus, the growth rate of shoots with air contact was about 3 times faster than the rate of fully submerged shoots when grown at air-equilibrium concentration of dissolved free CO2 in the water (16 [mu]M). This difference decreased with increased dissolved free CO2 concentration in the water, and the two shoot types grew at the same rate when the submerged shoots received >350 [mu]M free CO2. The quantitative importance of the floating rosette for total carbon uptake declined also with decreased ratio of floating rosette to total shoot weight. It is concluded that floating rosettes can enhance the inorganic carbon uptake of Callitriche. In contrast, air contact is of minor importance for nutrient transport.