Seasonal changes in temperature and nutrient control of photosynthesis, respiration and growth of natural phytoplankton communities

1. To investigate the influence of elevated temperatures and nutrients on photosynthesis, respiration and growth of natural phytoplankton assemblages, water was collected from a eutrophic lake in spring, summer, autumn, winter and the following spring and exposed to ambient temperature and ambient +2, +4 and +6 °C for 2 weeks with and without addition of extra inorganic nutrients. 2. Rates of photosynthesis, respiration and growth generally increased with temperature, but this effect was strongly enhanced by high nutrient availability, and therefore was most evident for nutrient amended cultures in seasons of low ambient nutrient availability. 3. Temperature stimulation of growth and metabolism was higher at low than high ambient temperature showing that long‐term temperature acclimation of the phytoplankton community before the experiments was of great importance for the measured rates. 4. Although we found distinct responses to relatively small temperature increases, the interaction between nutrient availability, time of the year and, thus, ambient temperature was responsible for most of the observed variability in phytoplankton growth, photosynthesis and respiration. 5. Although an increase in global temperature will influence production and degradation of organic material in lakes, the documented importance of ambient temperatures and nutrient conditions suggests that effects will be most pronounced during winter and early spring, while the remaining part of the growth season will be practically unaffected by increasing temperatures.     

Herbivory and growth in terrestrial and aquatic populations of amphibious stream plants

1. Many amphibious plant species grow in the transition between terrestrial and submerged vegetation in small lowland streams. We determined biomass development, leaf turnover rate and invertebrate herbivory during summer in terrestrial and aquatic populations of three amphibious species to evaluate advantages and disadvantages of aerial and submerged life.
2. Terrestrial populations had higher area shoot density, biomass and leaf production than aquatic populations, while leaf turnover rate and longevity were the same. Terrestrial populations experienced lower percentage grazing loss of leaf production (average 1.2–5.1%) than aquatic populations (2.9–17.3%), while the same plant dry mass was consumed per unit ground area.
3. Grazing loss increased linearly with leaf age apart from the youngest leaf stages. Grazing loss during the lifetime of leaves was therefore 2.4–3.1 times higher than mean apparent loss to standing leaves of all ages. The results imply that variation in density of grazers relative to plant production can account for differences in grazing impact between terrestrial and aquatic populations, and that fast leaf turnover keeps apparent grazing damage down.
4. We conclude that the ability of amphibious plants to grow submerged permits them to expand their niche and escape intense competition on land, but the stream does not provide a refugium against grazing and constrains plant production compared with the terrestrial habitat.     

Oxygen stress and reduced growth of Lobelia dortmanna in sandy lake sediments subject to organic enrichment

1. Lobelia dortmanna is a common representative of the small isoetid plants dominating the vegetation in nutrient‐poor lakes in Europe and North America. Because of large permeable root surfaces and continuous air lacunae Lobelia exchanges the majority of O₂ and CO₂ during photosynthesis across the roots. This leads to profound diel pulses of O₂ and CO₂ in sandy sediments with low microbial O₂ consumption rates. The ready radial root loss of O₂ may, however, make Lobelia very susceptible to more reducing sediments. Therefore, we grew Lobelia for 6 months on natural and organically enriched sandy sediments to test how: (i) root oxygenation influenced degradation of organic matter and depth profiles of N and C; (ii) Lobelia and microbial O₂ consumption rates influenced pool size and depth penetration of O₂ in the sediments; and (iii) sediment enrichment influenced growth and mineral nutrition of Lobelia. 2. Naturally low‐organic sediments (0.32% DW) accumulated organic C and N during the experiment as a result of growth of Lobelia and surface micro‐algae. In contrast, surface layers of enriched sediments (0.58, 0.87 and 2.46% DW) lost organic C and N because of enhanced mineralisation rates because of oxygen availability. In deeper layers of enriched sediments no significant differences in organic C and N pools were found between plant‐covered and plant‐free sediments probably because faster organic degradation because of root oxygenation was balanced by release of organic matter from the plants and because short roots with dense Fe‐Mn coatings in the most enriched sediments constrained O₂ release. 3. Depth‐integrated O₂ pools were much higher in light than darkness, higher in plant‐covered than plant‐free sediments and higher in sandy than in organically enriched sediments. All sediments had a primary O₂ maximum 1–2 mm below the sediment surface in light because of photosynthesis of micro‐algae. Plant‐covered sediments of low organic content (0.32 and 0.58% DW) also had a secondary deep maximum (2–4 cm) because of higher O₂ release from Lobelia roots than microbial O₂ consumption. Nitrification occurred here resulting in depletion of NH and accumulation of NO. In low organic sediments, oxygen pools increased with higher plant biomass both in light and darkness. The deep O₂ and NO₃ maxima disappeared in high organic sediments of greater O₂ consumption rates and smaller O₂ release rates. 4. Lobelia was stressed by increasing O₂ consumption rate of the sediments. Plant weight and leaf number declined twofold and maximum root length declined fourfold suggesting severe problems maintaining sufficient axial O₂ transport to the root tips because of rapid radial O₂ loss. Despite markedly higher nutrient concentrations in the enriched sediments, leaf‐N declined twofold and leaf‐P declined fourfold to growth‐limiting levels. These responses can be explained by constrains on mycorrhisal activity, root metabolism and vascular transport because of O₂ depletion. Management efforts to stop the decline and ensure the recovery of the isoetid vegetation should therefore focus on improving water quality as well as sediment suitability for growth.     

Variability of invertebrate herbivory on the submerged macrophyte Potamogeton perfoliatus

1. Invertebrate herbivory was studied in twenty-eight populations of the submerged macrophyte Potamogeton perfoliatus in Danish streams and lakes in mid-June. All populations but one experienced invertebrate herbivory and loss ranged from 0 to 11.9% of leaf area among populations. Loss generally increased with leaf age towards the base of the plants, and young apical leaves were rarely damaged. 2. Herbivory loss was significantly higher in streams (mean 5.0%) than in lakes (mean 2.2%), but varied greatly among populations within the same stream or lake and was not correlated to physico-chemical site characteristics, size or density of plant population, or leaf N and P content. High levels of invertebrate herbivory were therefore not associated with certain types of streams or lakes. 3. High herbivore biomass relative to abundance of plants was conducive to high loss. In streams, the biomass of the trichopteran Anabolia nervosa accounted for 50% of the variability in loss. No single species appeared to be equally important in lakes, although loss was correlated to the biomass of the chrysomelid beetle Macroplea appendiculata. Obligate herbivores, such as lepidopteran larvae, apparently exerted little damage on P. perfoliatus, and leaf mining and channelization from specialist feeders were negligible. It is concluded that shredders acting as facultative herbivores were the most important invertebrate herbivores on P. perfoliatus in Danish freshwaters.     

Photosynthetic Carbon Sources of Stream Macrophytes

Rates of photosynthesis of four submerged stream macrophyteswere examined under varying pH and composition of inorganiccarbon species. Callitriche stagnatis and Sparganium simplexused only CO₂ for photosynthesis. Potamogeton crispus and P.pectinatus used HCO₃ in addition to CO₂, but with much lowerefficiency. The photosynthetic rates at air equilibrium anda total inorganic carbon concentration of 5.0 mM were 2–3times lower than maximum rates at CO2 saturation for the HCO₃users and 10–14 times lower for the CO₂ users. The CO₂compensation point of entire plants of Callitriche (2.5 µM)and Sparganium (6.0µM) was well below the equilibriumconcentration (15 µM). and the low saturation points (250–500µM) also pointed to efficient use of CO2. Callitricheand Sparganium compete successfully with HCO₃^– users inhardwater streams, which have a higher exchange and generationcapacity of CO2 than stagnant and more soft waters. Rates ofphotosynthesis of Potamogeton crispus and P. pectinatus decreasedat high pH. Depending on the two alternative hypotheses forHCO₃use, this decline can be explained by CO₃^––inhibition of HCO₃^– uptake or by increasing capacity tobuffer H+efflux from the plant. Habitats subject to high pH,e. g. small ponds with dense vegetation, may have a strong selectionfor efficient mechanisms of HCO₃ use. Key words: Photosynthesis, Macrophytes, Carbon-source     

Patch dynamics of the stream macrophyte, Callitriche cophocarpa

1. We examined changes in position and growth of Callitriche cophocarpa patches in a shallow section of a small Danish stream during the main expansion of plant cover between April and June. Mean upstream growth of patches was only 7.5% of downstream growth. The mean growth rate was 1,02cm day^?1 in length and 0.55 cm day^?1 in width and the growth rates were not significantly dependent on patch size. For patches with an area (A) above 100cm², the growth rates in patch area (dA/dt) was proportional to patch circumference and thus √Aand the relative growth rate (dA/dt A^?1) was inversely proportional to √A.The smallest patches (<100cm²), however, expanded less than expected because of their combined tendency to grow more slowly in patch length and width than larger patches. 2. The expansion of plant cover will be much more rapid in many medium-sized patches compared to few large patches of a similar combined area, because internal selflimitation by light and space is partially relieved in smaller patches. We anticipate that recruitment and mortality of new-formed patches are critical steps preceding areal expansion. Quantification of these processes are needed fully to evaluate patch dynamics and space occupation.     

Rapid oxygen exchange across the leaves of Littorella uniflora provides tolerance to sediment anoxia

1. Littorella uniflora and Lobelia dortmanna are prominent small rosette species in nutrient‐poor, soft‐water lakes because of efficient root exchange of CO₂ and O₂. We hypothesise that higher gas exchange across the leaves of L. uniflora than of L. dortmanna ensures O₂ uptake from water and underlies its greater tolerance to sediment anoxia following organic enrichment. 2. We studied plant response to varying sediment O₂ demand and biogeochemistry by measuring photosynthesis, gas exchange across leaves and O₂ dynamics in plants during long‐term laboratory and field studies. Frequent non‐destructive sampling of sediment pore water was used to track changes in sediment biogeochemistry. 3. Addition of organic matter triggered O₂ depletion and accumulation of , Fe²⁺ and CO₂ in sediments. Gas exchange across leaf surfaces was 13–16 times higher for L. uniflora than for L. dortmanna. Oxygen in the leaf lacunae of L. uniflora remained above 10 kPa late at night on anoxic sediments despite organic enrichment. Leaf content of N and P of L. uniflora remained sufficient to keep up photosynthesis despite prolonged sediment anoxia, whereas nutrient content was too low for long‐term survival of L. dortmanna. 4. High gas exchange across L. uniflora leaves improves its performance and survival on anoxic sediments compared with L. dortmanna. Lobelia dortmanna uses the same gas‐tight leaves in air and water, which makes it highly susceptible to sediment anoxia but more cost‐effective in ultra‐oligotrophic environments because of slow leaf turnover.     

Vegetation and flow regime in lowland streams

1. The hydrological regime is important to the distribution of benthic organisms in streams. The objective of this study was to identify relationships between hydrological variables, describing the flow regime, and macrophyte cover, species richness, diversity and community composition in Danish lowland streams.
2. We quantified macrophyte vegetation in 44 Danish streams during summer by cover, species richness and diversity. Flow regime was characterized by 18 non-intercorrelated variables describing magnitude, frequency and duration of low and high flow events, timing or predictability of flow and general flow variability.
3. We found support in the stepwise multiple regressions analysis for our expectation that macrophyte cover is lowest in streams with high flow variability and highest in streams with long duration of low flow and low flow variability. We found support for the intermediate disturbance hypothesis as there were significant quadratic relationships between species richness and diversity as functions of disturbance frequency. There was poor discrimination in a detrended correspondence analysis (DCA) analysis of macrophyte community composition between four twinspan groups separating streams with different hydrological properties. Moreover, we did not find any relationship between the presence of disturbance-tolerant species and hydrological disturbance, suggesting that plant community composition developed independently of stream hydrology.     

Streamlining of plant patches in streams

1. Plants in shallow streams often grow in well‐defined monospecific patches experiencing a predictable unidirectional flow, though of temporally variable velocity. During maximum patch development in summer we studied: (i) the shape and streamlining of 59 patches of Callitriche cophocarpa, (ii) allometric relationships between canopy size and sediment area colonized by roots (root area) and (iii) fine‐scale flow gradients for a representative patch exposed to a range of velocities to evaluate relationships between patch shape and physical impact. 2. Canopy and root area viewed from above were elongated and streamlined in the flow direction, while uniform vegetative growth in all directions from a single colonizing shoot would have generated a circular form. Canopies were slightly wider in the upstream part than in the gradually tapering part downstream and the maximum height to length ratio averaged 0.25. The canopy and root area of the patches were more elongate and slender in sites with shallow water, where currents accelerate alongside patches and restrict lateral expansion, compared to deeper sites where currents can pass above the canopy. Similarly, the frontal area relative to planform area or root area was significantly lower in shallow water . Canopy shape and indices of streamlining did not change significantly with approach velocity (0.02–0.40 m s^?1), either because canopy shape is not sensitive to approach velocity or summer velocities were too low to induce such changes. 3. Sediment elevation within patches (avg. 4.1 cm) increased significantly with patch length, but did not differ between unstable sand or more stable coarse sediment for the same patch length. Shape of canopy and root area did not change significantly with sediment type. 4. Pressure drag on the canopy as a whole is probably reduced by its rounded front, restricted height and overall slender form with a low frontal area, while the downstream overhanging canopy increases drag compared to an ideal streamlined object. Across a 100‐fold range of root areas from 0.01 to 1 m², the frontal area of the canopy increased 29 times, planform area increased 38 times and volume increased 76 times, suggesting a trade‐off between physical impact of flow, light interception and anchoring strength. 5. The canopy was compressed at high approach velocities, with low current velocity within the canopy while steep velocity gradients developed across the exposed outer surfaces as the diverted flow accelerated. Because drag processes are additive, and exist at different spatial scales and Reynolds numbers on the surface and inside of plant canopies, direct measurements on entire canopies under controlled conditions are needed to test the functional importance of their shape, size and porosity to flow.     

Influence of sediment organic enrichment and water alkalinity on growth of aquatic isoetid and elodeid plants

1. Lake eutrophication has increased phytoplankton blooms and sediment organic matter. Among higher plants, small, oligotrophic rosette species (isoetids) have disappeared, while a few tall, eutrophic species (elodeids) may have persisted. Despite recent reduction of nutrient loading in restored lakes, the vegetation has rarely regained its former composition and coverage. Patterns of recovery may depend on local alkalinity because HCO₃^? stimulates photosynthesis of elodeids and not of isoetids. In laboratory growth experiments with two isoetids (Lobelia dortmanna and Littorella uniflora) and two elodeids (Potamogeton crispus and P. perfoliatus), we test whether organic enrichment of lake sediments has a long‐lasting influence by: (i) reducing plant growth because of oxygen stress on plant roots and (ii) inhibiting growth more for isoetids than elodeids. We also test whether (iii) increasing alkalinity (from 0.17 to 3.20 meq. L^?1) enhances growth and reduces inhibition of organic sediment enrichment for elodeids but not for isoetids. 2. In low organic sediments, higher oxygen release from roots of isoetids than elodeids generated oxic conditions to greater sediment depth for Lobelia (4.3 cm) and Littorella (3.0 cm) than for Potamogeton species (1.6–2.2 cm). Sediment oxygen penetration depth fell rapidly to 0.4–1.0 cm for all four species at even modest organic enrichment and oxygen consumption in the sediments. Roots became shorter and isoetid roots became thicker to better supply oxygen to apical meristems. 3. Growth of elodeids was strongly inhibited across all levels of organic enrichment of sediments being eight‐fold lower at the highest enrichment compared to the unenriched control. Leaf biomass of isoetids increased three‐fold by moderate organic enrichment presumably because of greater CO₂ supply from sediments being their main CO₂ source. At higher organic enrichment, isoetid biomass was reduced, leaf chlorophyll declined up to 10‐fold, root length declined from 7 to <2 cm and mortality rose (up to 50%) signalling high plant stress. 4. Lobelia was not affected by HCO₃^? addition in accordance with its use of sediment CO₂. Biomass of elodeids increased severalfold by rising alkalinity from 0.17 to 3.20 meq. L^?1 in accordance with their use of HCO₃^? for photosynthesis, while the negative impact of organically enriched sediments remained. 5. Overall, root development of all four species was so strongly restricted in sediments enriched with labile organic matter that plants if growing in situ may lose root anchorage. Other experiments demonstrate that this risk is enhanced by greater water content and reduced consolidation in organically rich sediments. Therefore, formation of more muddy and oxygen‐demanding sediments during eutrophication will impede plant recovery in restored lakes while high local alkalinity will help elodeid recovery.