Wilhelm Granéli

THE INFLUENCE OF MULTIPLE INTRODUCED PREDATORS ON A LITTORAL POND COMMUNITY

In a replicated field experiment we studied the effects of natural densities of two exotic consumers, the predatory and herbivorous signal crayfish (Pacifastacus leniusculus) and the predatory rainbow trout (Oncorhynchus mykiss), on multiple trophic levels of a pond community. The goals were to: (1) determine the individual and combined effects of predators on macroinvertebrates, macrophytes, and periphytic algae; (2) evaluate the strength of direct and indirect interactions in a food web influenced by omnivores; and (3) evaluate the relative importance of direct and indirect predator effects on mortality and growth of a native frog species, Rana temporaria. The experiment showed that both signal crayfish and rainbow trout had strong effects on multitrophic levels of a littoral pond community, through direct consumption and indirect effects on lower trophic levels. Crayfish had weak but significant negative effects on the biomass of predatory invertebrates and greatly reduced the biomass of snails, the most abundant invertebrate grazers. Although the number of active herbivorous tadpoles tended to be higher in crayfish cages relative to control cages, the proportion of surviving froglets was lower in crayfish cages than in control cages, possibly due to crayfish predation on injured tadpoles. The size of surviving froglets did not differ from controls, but tadpoles in crayfish cages often suffered tail injuries. Macrophyte coverage decreased as a result of crayfish consumption and nonconsumptive fragmentation. However, the biomass of periphyton increased in crayfish cages relative to controls, probably due to reduced grazing from snails. In contrast, trout had strong negative effects on the biomass of both predatory invertebrates and insect grazers, whereas trout had less effect on snail biomass than did crayfish. Also, in contrast to crayfish cages, the number of active tadpoles in trout cages was lower than in controls, probably due to a combination of trout predation and trout-induced reduced tadpole activity. Trout had a strong negative impact on froglet survival, and froglets in trout cages metamorphosed at a smaller size and had reduced growth rates compared to froglets in crayfish and control cages. As with crayfish, the biomass of periphyton increased in trout cages relative to controls, which may be due to a combination of both density and trait-mediated trout effects on tadpole grazing. In treatments with multiple predators the effects of crayfish and trout on caged communities were independent, and there were few interactions. Mostly effects of combined predators reflected those in single predator cages. Our results demonstrate that noninteracting, introduced multiple predators can have strong direct and indirect effects on multiple trophic levels in pond communities. Trophic cascades may develop in aquatic food webs even with omnivores such as crayfish, and in complex habitats with trout. These strong indirect effects are mediated through both predation on important grazers (i.e., the crayfish-snail-periphyton link) and a combination of density and behavioral responses of grazers to predators (i.e., the trout-tadpole-periphyton link). When two noninteracting predators have strong but different effects on prey survival or activity, their combined effects on intermediate trophic levels reflect responses to the more dangerous predator. (Less)

Influence of Aquatic Macrophytes on Phosphorus Cycling in Lakes

Emergent macrophytes take up their phosphorus exclusively from the sediment. Submerged species obtain phosphorus both from the surrounding water and from the substrate, but under normal pore and lake water phosphorus concentrations, substrate uptake dominates. Release of phosphorus from actively growing macrophytes (both submerged and emergent) is minimal and epiphytes obtain phosphorus mainly from the water. Decaying macrophytes may act as an internal phosphorus source for the lake and add considerable quantities of phosphorus to the water. A large part of the released phosphorus is often retained by the sediments. In perennial macrophytes the amount of phosphorus released from decaying shoots is dependent on the degree of phosphorus conservation within the plant. Macrophyte stands may also be a permanent phosphorus sink due to burial of plant litter. Macrophytes affect the chemical environment (oxygen, pH), which in turn has effects on the phosphorus cycling in lakes. However, the impact of aquatic macrophytes on whole-lake phosphorus cycling is largely unknown. Controlled full-scale harvesting, herbicide or herbivory experiments are almost totally lacking. Emergent macrophytes respond positively to eutrophication, but fertilization experiments have shown that nitrogen rather than phosphorus may be the key element. Submerged macrophytes are adversely affected by a large increase in the external phosphorus input to a lake. This effect may be caused by epiphyte shading, phytoplankton shading or deposition of unfavourable sediments.

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)

Influence of an Exotic and a Native Crayfish Species on a Littoral Benthic Community

The aim of this study was to compare the effects of the introduced signal crayfish (Pacifastacus leniusculus) and the native noble crayfish (Astacus astacus) on a benthic food web. We mimicked the habitat of a pond littoral in 4.5-m 2 plastic pools stocked with natural densities of macrophytes, invertebrates and either signal crayfish, noble crayfish, or kept as crayfish free controls. After two summer months, all invertebrates and macrophytes were collected from each pool, and periphyton was sampled on one substratum exposed and two substrata not exposed to crayfish grazing. Samples for stabile isotope analysis of benthos were collected in pools with noble crayfish. 15 N ratios showed that crayfish were top consumers, and 13 C ratios indicated that they received most of their carbon from invertebrates, but less from primary producers. Crayfish did not affect the biomass of predatory invertebrates, dominated by active swimmers among Heteroptera and Coleoptera, but had a strong impact on grazers dominated by thin-shelled Lymmaea snails. Hard-shelled Bithynia snails were also reduced in numbers, but the largest of these snails were consumed less than thin-shelled Lymmaea snails. The reduced biomass of snails had an indirect positive effect on periphyton biomass on all three substrata. Crayfish grazed selectively on macrophytes and reduced the biomass of Chara, whereas Elodea was less affected. The exotic signal crayfish had, overall, a stronger impact on the biomass of macrophytes and grazers than the native noble crayfish. The results indicate that crayfish may structure food webs through consumption from many food levels. The short-term influence of crayfish on other trophic levels depends on crayfish feeding efficiency, food preferences and species-specific consumption rates.

The influence of animals on phosphorus cycling in lake ecosystems

Aquatic animals directly influence the cycling of phosphorus in lakes through feeding and excretion. Traditionally, animals (zooplankton, benthic invertebrates and fish) have been assigned only minor roles in the process of freshwater phosphorus cycling. They were regarded as consumers without much regulating influence. Today there is growing evidence that animals, predators and herbivores, directly or indirectly can control biomass of primary producers and internal cycling of phosphorus.This paper summarizes different mechanisms of transformation and translocation of phosphorus via different groups of organisms.

Effect of Humic Substance Photodegradation on Bacterial Growth and Respiration in Lake Water

ABSTRACT This study addresses how humic substance (HS) chemical composition and photoreactivity affect bacterial growth, respiration, and growth efficiency (BGE) in lake water. Aqueous solutions of HSs from diverse aquatic environments representing different dissolved organic matter sources (autochthonous and allochthonous) were exposed to artificial solar UV radiation. These solutions were added to lake water passed through a 0.7-μm-pore-size filter (containing grazer-free lake bacteria) followed by dark incubation for 5, 43, and 65 h. For the 5-h incubation, several irradiated HSs inhibited bacterial carbon production (BCP) and this inhibition was highly correlated with H2O2 photoproduction. The H2O2 decayed in the dark, and after 43 h, nearly all irradiated HSs enhanced BCP (average 39% increase relative to nonirradiated controls, standard error = 7.5%, n = 16). UV exposure of HSs also increased bacterial respiration (by ∼18%, standard error = 5%, n = 4), but less than BCP, resulting in an average increase in BGE of 32% (standard error = 10%, n = 4). Photoenhancement of BCP did not correlate to HS bulk properties (i.e., elemental and chemical composition). However, when the photoenhancement of BCP was normalized to absorbance, several trends with HS origin and extraction method emerged. Absorbance-normalized hydrophilic acid and humic acid samples showed greater enhancement of BCP than hydrophobic acid and fulvic acid samples. Furthermore, absorbance-normalized autochthonous samples showed ∼10-fold greater enhancement of BCP than allochthonous-dominated samples, indicating that the former are more efficient photoproducers of biological substrates.

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.

Photoproduction of dissolved inorganic carbon in temperate and tropical lakes – dependence on wavelength band and dissolved organic carbon concentration

We have evaluated photoeffects of UV-B, UV-A and PAR radiation on dissolved organic matter (DOM). Photochemical production of dissolved inorganic carbon (DIC) was measured in sterile lake water from Sweden and Brazil after 6 hours of sun exposure. Tubes were exposed to four solar radiation regimes: Full-radiation, Full-radiation minus UV-B, Full-radiation minus UV-B and UV-A (PAR) and darkness.In both areas, lakes with most DOC (varying between 3 and 40 mg C l-1) were highly humic, resulting in high UV-B attenuation coefficients (Kd = 5–466 m-1). Under Full-radiation, photooxidative DIC-production varied from 0.09 to 1.7 mg C l-1per 6 h, without UV-B from 0.07 to 1.4 mg C l-1 and with PAR only from 0.02 to 0.7 mg C l-1. UV-B radiation explains a minor part (17%) of the photoooxidative DIC-production, while UV-A and PAR have larger effects (39% and 44%, respectively). Photooxidation was proportional to DOC-content and DIC-production was positively related to decrease in DOC and to loss of absorbance at 250 nm. There was no significant difference in DOC and radiation normalized DIC-production between Swedish and Brazilian lakes. The UV-B dose during incubations was approximately 3 times higher in Brazil compared to Sweden, while UV-A and PAR doses were similar. We conclude that DOC from tropical and temperate freshwaters do not seem to differ with respect to sensitivity to photooxidation.

Internal phosphorus loading in Lake Ringsjön

Lake Ringsjon did not respond with decreased algal production following a substantial reduction in external phosphorus loading. This is typical of many shallow lakes which for decades have received excessive amounts of nutrients. The inertia is due to large amounts of phosphorus (P) stored in sediments and biota, causing internal phosphorus loading. Much of this phosphorus is thought to be released from the organic-rich profundal sediments. In Lake Ringsjon, only one third of the total bottom area is covered by such sediments, the rest being dominated by sand and silt. In the profundal sediments bulk P content was not exceptionally high (approximately 2 mg P : g DW 1 ), while the pore water phosphate concentrations, especially in Satofta Basin, were very high, indicating large potential for phosphorus release to the water. This is also indicated by the large proportion of Fe- and Albound P in the sediments of Satofta Basin. Although there are no direct quantifications of phosphorus release from the sediments in Lake Ringsjon, measurements of phosphorus concentrations in the water mass as well as budget calculations for the three basins clearly show a high capacity for internal loading. Phosphorus concentrations generally increase during summer, when external additions are minimal. Until 1980, the annual external phosphorus addition to Lake Ringsjon greatly exceeded the output, showing that the lake was an efficient phosphorus trap. Since then, input and output have been balanced, but in recent years signs that the lake is once again retaining phosphorus on an annual basis are evident. There are marked differences between the three basins, with Western Basin generally retaining phosphorus, while the upstream Eastern Basin and Satofta Basin during the 1980s often exported phosphorus. It is not possible to evaluate the effects of the fish biomanipulation on the internal loading of phosphorus from the sediment.

Bacterial growth on allochthonous carbon in humic and nutrient-enriched lakes: Results from whole-lake C-13 addition experiments

Organic carbon (C) in lakes originates from two distinct sources—primary production from within the lake itself (autochthonous supply) and importation of organic matter from the terrestrial watershed (allochthonous supply). By manipulating the 13C of dissolved inorganic C, thereby labeling within-lake primary production, we examined the relative importance of autochthonous and allochthonous C in supporting bacterial production. For 35 days, NaH13CO3 was added daily to two small, forested lakes. One of the lakes (Peter) was fertilized so that primary production exceeded total respiration in the epilimnion. The other lake (Tuesday), in contrast, was low in productivity and had high levels of colored dissolved organic C (DOC). To obtain bacterial C isotopes, bacteria were regrown in situ in particle-free lake water in dialysis tubes. The contribution of allochthonous C to bacterial biomass was calculated by applying a two-member mixing model. In the absence of a direct measurement, the isotopic signature of the autochthonous end-member was estimated indirectly by three different approaches. Although there was excess primary production in Peter Lake, bacterial biomass consisted of 43–46% allochthonous C. In Tuesday Lake more than 75% of bacterial growth was supported by allochthonous C. Although bacteria used autochthonous C preferentially over allochthonous C, DOC from the watershed contributed significantly to bacterial production. In combination with results from similar experiments in different lakes, our findings suggest that the contribution of allochthonous C to bacterial production can be predicted from ratios of chromophoric dissolved organic matter (a surrogate for allochthonous supply) and chlorophyll a (a surrogate for autochthonous supply).