Stefan E.B. Weisner

Phenotypic plasticity in Phragmites australis as a functional response to water depth

We have performed investigations to see if the emergent macrophyte Phragmites australis (Cav.) Trin. ex Steud. exhibits phenotypic plasticity as a response to water depth and if such responses in biomass allocation pattern and morphology are functional responses, improving the performance of the plant. In greenhouse experiments plants were grown in deep or shallow water to evaluate plastic responses. Allometric methods were used to handle effects caused by size differences between treatments. To evaluate if phenotypic responses to water depth are functional, the relative growth rate (RGR) of plants acclimatised to shallow or deep water, respectively, were compared in deep water, and the growth of plants in fluctuating and constant water level were compared. When grown in deep (70 or 75 cm), compared to shallow (20 or 5 cm) water, plants allocated proportionally less to below-ground weight, made proportionally fewer but taller stems, and had rhizomes that were situated more superficially in the substrate. Plants acclimatised to shallow water had lower RGR than plants acclimatised to deep water, when they were grown in deep water, and plants in constant water depth (40 cm) grew faster than plants in fluctuating water depth (15/65 cm). In an additional field study, the rhizomes were situated superficially in the sediment in deep, compared to shallow water. We have shown that P. australis acclimatises to deep water with phenotypic plasticity through allocating more resources to stem weight, and also by producing fewer but taller stems, which will act to maintain a positive carbon balance and an effective gas exchange between aerial and below-ground parts. Furthermore, the decreased proportional allocation to below-ground parts probably results in decreased nutrient absorption, decreased anchorage in the sediment and decreased carbohydrate reserves. Thus, in deep water, plants have an increased risk of becoming uprooted and experience decreased growth and dispersal rates. (Less)

Mechanisms regulating abundance of submerged vegetation in shallow eutrophic lakes

Abstract Shallow eutrophic lakes tend to be either in a turbid state dominated by phytoplankton or in a clear-water state dominated by submerged macrovegetation. Recent studies suggest that the low water turbidity in the clear-water state is maintained through direct and indirect effects of the submerged vegetation. This study examined what mechanisms may cause a recession of the submerged vegetation in the clear-water state, and thereby a switch to the turbid state. The spatial distribution of submerged vegetation biomass was investigated in two shallow eutrophic lakes in the clear-water state in southern Sweden. Biomass of submerged vegetation was positively correlated with water depth and wave exposure, which also were mutually correlated, suggesting that mechanisms hampering submerged vegetation were strongest at shallow and/or sheltered locations. The growth of Myriophyllum spicatum, planted in the same substrate and at the same water depth, was compared between sheltered and wave exposed sites in two lakes. After 6 weeks the plants were significantly smaller at the sheltered sites, where periphyton production was about 5 times higher than at the exposed sites. Exclosure experiments were conducted to evaluate the effects of waterfowl grazing on macrophyte biomass. Potamogeton pectinatus growth was decreased by grazing, whereas M. spicatum was not affected. The effects were greater at a sheltered than at a wave-exposed site, and also negatively related to distance from the reed belt. These results suggest that competition from epiphytes and waterfowl grazing hamper the development of submerged vegetation at sheltered and/or shallow locations. An increased strength of these mechanisms may cause a recession of submerged vegetation in shallow eutrophic lakes in the clear-water state and thereby a switch to the turbid state.

Rhizome dynamics and resource storage in Phragmites australis

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

Indirect effects of fish community structure on submerged vegetation in shallow, eutrophic lakes: an alternative mechanism

The loss of submerged macrophytes during eutrophication of shallow lakes is a commonly observed phenomenon. The proximate reason for this decline is a reduction of available light due to increasing phytoplankton and/or epiphyton biomass. Here we argue that the ultimate cause for the transition from a macrophyte-dominated state to a phytoplankton-dominated state is a change in fish community structure. A catastrophic disturbance event (e.g. winterkill) acting selectively on piscivores, cascades down food chains, eventually reducing macrophyte growth through shading by epiphyton, an effect that is reinforced by increasing phytoplankton biomass. The transition back from the phytoplankton to the macrophyte state depends on an increase in piscivore standing stock and a reduction of planktivores. A conceptual model of these mechanisms is presented and supported by literature data and preliminary observations from a field experiment.

Influence of substrate conditions on the growth of Phragmites australis after a reduction in oxygen transport to below-ground parts

Abstract The effect of a reduction in O 2 transport through culms to below-ground parts was compared between two monospecific stands of Phragmites australis (Cav.) Trin. ex Steudel growing in sand and calcareous mud (redox potentials about 220 mV and −10 mV in the upper 30 cm, respectively). Oxygen transport was reduced by cutting the culms above and below the water surface. The effects from cutting in June, when shoot growth is rapid and carbohydrate supplies in the rhizomes are at a minimum, was compared with cutting in August, when shoot growth is slower and rhizomes are recharged with carbohydrates. Cutting the reed below water in June, in the stand growing in mud, almost totally inhibited the regrowth of shoots the following summer, while cutting Above water reduced regrowth of shoots. Cutting in August did not, however, reduce growth the following summer. The growth, the following summer, of the stand growing in sand was not affected by any cutting treatments. Oxygen concentration in rhizomes decreased more after cutting below water on the muddy site than on the sandy site. Oxygen supply was also more affected by cutting below water than above water. The need for O 2 in below-ground parts may be higher in the more reducing substrate, where O 2 diffuses from the roots and is consumed in the substrate or by reducing substances diffusing into roots and rhizomes. Excessive eutrophication may thus affect the reed negatively, through the deposition of highly reducing sediments. Inhibition of O 2 transport to below-ground parts by shoot grazing or increased water depth may have a detrimental effect on reeds growing in reducing substrates.

Rhizome architecture inPhragmites australis in relation to water depth: Implications for within-plant oxygen transport distances

Phragmites australis (Cav.)Trin. exSteud. is a perennial plant, largely relying on its rhizomes for resource storage, spreading and anchorage in the substrate. Vertical distribution and length of horizontal rhizomes ofPhragmites australis were investigated at the reed bed edge in a lake in southern Sweden. In deep water, horizontal rhizomes were relatively short and superficially situated in the substrate. It is hypothesised that this is an adaptation to water depth by keeping O2-transport distances through shoots and rhizomes as short as possible. In shallow water,P. australis rhizomes generally penetrated deeply into the substrate, probably improving anchorage and nutrient uptake possibilities. Further, horizontal rhizomes were longer in shallow water, which may increase the rate of vegetative spread. Because of these changes in rhizome architecture, “critical within-plant oxygen transport distances” did not change with water depth. This indicates thatP. australis maximises the extension of its rhizomes in relation to spatial differences in water depth. This may limit the ability ofP. australis to tolerate sudden temporal increases in water depth or eutrophication.

Factors affecting the internal oxygen supply of Phragmites australis (Cav.) Trin. ex. Steudel in situ

Abstract Oxygen concentration in basal stems of Phragmites australis (Cav.) Trin. ex Steudel was measured in situ, in monospecific stands. The mean O 2 concentration in basal stems, and mean shoot length above water, decreased in deeper water. Oxygen concentration in basal stems was correlated to shoot length above water at a given water depth, and the mean O 2 concentration in basal stems of shoots with similar length above water decreased in deeper water. The results suggest that O 2 transport from shoots to below-ground parts is restricted in deep water by: (1) small shoot length above water; (2) long O 2 transport distance. No influence of substrate redox potential on O 2 concentration in basal stems was found.

Influence of germination time on juvenile performance of Phragmites australis on . temporarily exposed bottoms-implications for the colonization of lake beds

Three cohorts of seedlings of Phragmites australis (Cav.) Trin ex Steud., germinated in May, June and July, were allowed to grow in shallow water (depth 5 cm or less) in southern Sweden. In the autumn, size parameters were measured on the plants. In the second year, the water level was raised to 0.8 m and emergence of shoots, plant survival and size parameters were recorded. The mean plant weight by the end of the first year differed markedly between cohorts. Rhizome biomass showed a relationship of 700:70:1 between the May, June and July cohorts. In the second year, rate of emergence above the water surface, and maximum height of plants that did not reach the water surface, was positively related to the size (mass) the plants had achieved after the first year. Only plants that emerged above the water surface survived the second summer, resulting in survival rates for the May, June and July cohorts of 90%, 68% and 0%, respectively. The rhizome weight of the smallest survivors had decreased after the second summer compared with values after the first summer. Hence, they were not capable of ‘reloading’ their rhizomes during the second year. In a temperate climate, the size of juvenile plants after the first year, which is strongly dependent on early germination on exposed bottoms (i.e. bottoms without standing water), determines their water depth tolerance during the second year. The timing and duration of exposure, as well as the subsequent depth of re-flooding, are all of fundamental importance for successful ‘lakeward’ seedling expansion of P. australis.

Decoupling of cascading trophic interactions in a freshwater, benthic food chain

Food chain theory provides explicit predictions for equilibrium biomasses among trophic levels in food chains of different lengths. Empirical studies on freshwater benthic food chains have typically been performed on chains with up to three levels and in field experiments with limited spatial and temporal scale. Here we use a “natural snapshot experiment” approach to study equilibrium biomass and abundance among trophic levels in natural ponds differing only with respect to fish assemblage structure. Forty-four ponds were surveyed for their densityand biomass of fish, snails and periphyton. Ponds were divided into three categories based on fish assemblage: ponds with no fish (two trophic levels), ponds with molluscivorous fish (three trophic levels) and ponds that also had piscivorous fish (four trophic levels). Ponds without fish had a high density and biomass of snails and a low biomass of periphyton, whereas snails were scarce and periphyton biomass was high in ponds with molluscivorous fish. In the presence of piscivores, molluscivore populations consisted of low numbers of large individuals. Snail assemblages in piscivore ponds were characterised by relativelyhigh densities of small-bodied detritivorous species and periphyton biomass was not significantlydifferent from ponds with three trophic levels. Thus, predictions from classic food chain theory were upheld in ponds with up to three trophic levels. In ponds with four trophic levels, however, there was a decoupling of the trophic cascade at the piscivore-molluscivore level. Gape-limited piscivory, predation on snails by molluscivores that have reached an absolute size refuge from predation, and changes in food preferences of the dominant snails are suggested to explain the observed patterns.

Wave exposure related growth of epiphyton : implications for the distribution of submerged macrophytes in eutrophic lakes

The distribution of submerged macrophytes in eutrophic lakes has been found to be skewed towards sites with intermediate exposure to waves. Low submerged macrophyte biomass at exposed sites has been explained by, for instance, physical damage from waves. The aim of this study was to investigate if lower biomass at sheltered sites compared to sites with intermediate exposure to waves can be caused by competition from epiphyton.Investigations were performed in eutrophic lakes in southern Sweden. Samples of submerged macrophytes and epiphytic algae on the macrophytes were taken along a wave exposure gradient. The amount of epiphyton (AFDW) per macrophyte biomass decreased with increased exposure. Biomass of submerged macrophytes, on the other hand, increased with increased exposure until a relatively abrupt disappearance of submerged vegetation occurred at high exposures. Production of epiphytic algae was monitored on artificial substrates from June to September at a sheltered and an exposed site in three lakes. It was higher at sheltered sites compared with exposed sites.We suggest that epiphytic algae may be an important factor in limiting the distribution of submerged macrophytes at sheltered sites in eutrophic lakes.