Cadmium tolerance and adsorption by the marine brown alga Fucus vesiculosus from the Irish Sea and the Bothnian Sea

Cadmium (Cd) uptake capacities and Cd tolerance of the marine alga Fucus vesiculosus from the Irish Sea (salinity 35 psu) and from the Bothnian Sea (northern Baltic, 5 psu) were quantified. These data were complemented by measurements of changes in maximal photosynthetic rate (P(max)), dark respiration rate and variable fluorescence vs. maximal fluorescence (F(v):F(m)). At concentrations between 0.01 and 1 mmol Cd l(-1), F. vesiculosus from the Bothnian Sea adsorbed significantly more (about 98%) Cd compared with F. vesiculosus from the Irish Sea. The photosynthetic measurements showed that the Bothnian Sea F. vesiculosus were more sensitive to Cd exposure than the Irish Sea algae. The algae from the Irish Sea showed negative photosynthetic effects only at 1 mmol Cd l(-1), which was expressed as a decreased P(max) (-12.3%) and F(v):F(m) (-4.6%). On the contrary, the algae from the Bothnian Sea were negatively affected already at Cd concentrations as low at 0.1 mmol Cd l(-1). They exhibited increased dark respiration (+11.1%) and decreased F(v):F(m) (-13.9%). The results show that F. vesiculosus from the Bothnian Sea may be an efficient sorption substrate for Cd removal from Cd contaminated seawater and this algae type may also have applications for wastewater treatment.     

Intriguing asexual life in marginal populations of the brown seaweed Fucus vesiculosus

Reproduction of attached large brown algae is known to occur only by sexual zygotes. Using microsatellites we show evolution of asexual reproduction in the bladder wrack promoting population persistence in the brackish water Baltic Sea (< 6 psu). Here a dwarf morph of Fucus vesiculosus is dominated by a single clone but clonal reproduction is also present in the common form of the species. We describe a possible mechanism for vegetative reproduction of attached algae, and conclude that clonality plays an important role in persistence and dispersal of these marginal populations, in which sexual reproduction is impaired by low salinity.     

REPRODUCTIVE SUCCESS OF FUCUS VESICULOSUS (PHAEOPHYCEAE) IN THE BALTIC SEA

Marine organisms colonizing brackish habitats such as the Baltic Sea must cope with the negative effects of low salinities on reproductive success because these may reduce gamete viability and/or increase polyspermy. Reproductive characteristics of the marine seaweed Fucus vesiculosus L. were studied in several brackish habitats, particularly in the northern Baltic Sea, to understand its ability to reproduce where few other marine species survive. Polyspermy and fertilization success were variable at the boundary of the continuous distribution of F. vesiculosus in the Baltic Sea, and polyspermy was high (10%–30%) when fertilization was successful. A strong female bias (80%–86%, ca. 5.5:1) was found at the northernmost limit of Baltic F. vesiculosus. Electrophysiological studies showed that many eggs have a high input resistance (519 ± 150 MΩ[mean ± SE, n = 14] at Drivan, 1995), which may be helpful in preventing polyspermy in this brackish habitat. The polyspermy block remains sodium-dependent in the northern Baltic. Sperm bound quickly to northern Baltic eggs in natural water, but fertilization was delayed compared to marine F. vesiculosus. A subset of northern Baltic eggs studied during an optimal reproductive period (7–11 July 1995) had a membrane potential (E_m) of ca. −100 mV and an effective fertilization potential (FP) of ca. 2 min with a plateau of −25 mV, but repolarized too rapidly for the FP to be protective. Pronuclear migration and cell wall secretion occurred more slowly in Baltic than in marine zygotes. The reproductive success of theseboundary populations may be dependent upon windows of opportunity when there are favorable combinations of the levels of salinity, water motion, population density, and sex ratio. These factors and the short duration of the reproductive season in the northern Baltic Sea may result in reproductive failure in some years.     

FLUORESCENCE EMISSION SPECTRA OF MARINE AND BRACKISH‐WATER ECOTYPES OF FUCUS VESICULOSUS AND FUCUS RADICANS (PHAEOPHYCEAE) REVEAL DIFFERENCES IN LIGHT‐HARVESTING APPARATUS1

The Bothnian Sea in the northerly part of the Baltic Sea is a geologically recent brackish‐water environment, and rapid speciation is occurring in the algal community of the Bothnian Sea. We measured low‐temperature fluorescence emission spectra from the Bothnian Sea and the Norwegian Sea ecotypes of Fucus vesiculosus L., a marine macroalga widespread in the Bothnian Sea. Powdered, frozen thallus was used to obtain undistorted emission spectra. The spectra were compared with spectra measured from the newly identified species Fucus radicans Bergström et L. Kautsky, which is a close relative of F. vesiculosus and endemic to the Bothnian Sea. The spectrum of variable fluorescence was used to identify fluorescence peaks originating in PSI and PSII in this chl c–containing alga. The spectra revealed much higher PSII emission, compared to PSI emission, in the Bothnian Sea ecotype of F. vesiculosus than in F. radicans or in the Norwegian Sea ecotype of F. vesiculosus. The results suggest that more light‐harvesting chl a/c proteins serve PSII in the Bothnian Sea ecotype of F. vesiculosus than in the two other algal strains. Treatment of the Bothnian Sea ecotype of F. vesiculosus in high salinity (10, 20, and 35 practical salinity units) for 1 week did not lead to spectral changes, indicating that the measured features of the Bothnian Sea F. vesiculosus are stable and not simply a direct result of exposure to low salinity.     

LOCAL ADAPTATION OF CERAMIUM TENUICORNE (CERAMIALES,RHODOPHYTA) WITHIN THE BALTIC SEA1

Ceramium tenuicorne (Kützing) Wærn is a red alga that is widely distributed in the brackish Baltic Sea. We studied the growth response of Ceramium to low salinity and nutrient enrichment, using isolades from two regions of the Baltic Sea where the alga approaches its inner distribution limit. Ecotypic differentiation was observed in that differences in growth response among isolates corresponded to salinity conditions in their regions of origin. Isolates from the Gulf of Bothnia (4 psu) had inherently lower growth rates that were not increased when transferred to higher salinity, but were better adapted to very low salinity levels than isolates from the Baltic Proper (7 psu). Further, the results indicate that Ceramium from different regions of the Baltic Sea vary in their responses to nutrient enrichment. The observed differences may be best described as a quantitative difference in the proportion of isolates with hyposaline adaptation. The results indicate that the wide distribution of Ceramium in the Baltic Sea is better explained by the occurrence of locally adapted genotypes than by a generalist life strategy, and provide example of adaptive differentiation in a marine edge environment.     

Photosynthesis in relation to D1, PsaA and Rubisco in marine and brackish water ecotypes of Fucus vesiculosus and Fucus radicans (Phaeophyceae)

The aim of this study was to investigate photosynthetic differences between the marine, Norwegian Sea ecotype and the brackish, Bothnian Sea ecotype of F. vesiculosus and F. radicans and to see whether photosynthetic differences could be connected with the relative amounts of D1 protein (PSII), PsaA (PSI) protein and/or Rubisco. For this purpose, we tested if a higher photosynthetic maximum (P _max) in the Atlantic Ocean ecotype of F. vesiculosus relative to the Baltic Sea ecotype, and an increase of the P _max in Baltic Sea ecotype of F. vesiculosus at higher salinity, could be due to an increase in the relative amounts of Rubisco. The proteins have been evaluated on a relative basis. Immunoblot signals showed that the amount of Rubisco was higher in both ecotypes of F. vesiculosus than in F. radicans, but no differences could be detected between the two ecotypes of F. vesiculosus. The results suggest an uneven photosystem protein stoichiometry in Fucus, with more of the PSI protein PsaA relative to the PSII protein D1. The difference in P _max between the two ecotypes of F. vesiculosus might be related to the difficulties for the algae to adapt to the environment in Bothnian Sea.     

Despite marine traits,the endemic Fucus radicans (Phaeophyceae) is restricted to the brackish Baltic Sea

Many of the marine species that were introduced to the Baltic Sea during the Littorina stage (c. 8500–3000 years BP), e.g. Fucus vesiculosus and F. serratus, have adapted to the present low salinity. These marine species have gone from marine conditions into lower salinity environments. In this paper we ask why the recently discovered endemic brown alga Fucus radicans shows the opposite pattern. Fucus radicans is only present in the northern parts of the Baltic Sea, the low salinity Bothnian Sea (4–6 psu). Potentially, the fitness of F. radicans might be reduced in higher salinities if it is better adapted to brackish conditions. We hypothesize, however, that the southern distribution limit of F. radicans is set by biotic factors, e.g. competition with F. vesiculosus and higher grazing pressure by Idotea balthica and not by salinity. Our results show that the reproductive output of F. radicans is limited by low salinity (4 psu) but increases in higher salinities. However, the southern distribution limit, i.e. the northern Baltic Proper, is regulated by biotic factors, where the additive effects from shading by taller F. vesiculosus thalli and grazing on F. radicans by the isopod I. balthica limit the biomass production of F. radicans. We suggest that F. radicans still maintains marine traits due to its ability to propagate clonally and is restricted to the Bothnian Sea by interactions with F. vesiculosus and I. balthica. We also propose that increased precipitation due to climate change might affect the northern range limit and that the distribution of F. radicans could be expected to shift further south into the Baltic Proper.     

Osmotic and ionic haemolymph concentrations in the Baltic Sea amphipod Gammarus oceanicus in relation to water salinity

The aim of this work was to determine the osmotic and ionic (Na(+), K(+), Ca(2+), Mg(2+) and Cl(-)) haemolymph concentrations in Gammarus oceanicus at different salinity levels. Being a species of marine origin it inhabits brackish waters of the Baltic Sea. G. oceanicus specimens were collected in January 2003 from the Gulf of Gdansk (salinity 7 psu). The animals were gradually acclimated to eight different salinity levels (5, 7, 14, 20, 25, 30, 35 and 41 psu) at a temperature of 5 degrees C and 100% oxygen saturation. The haemolymph osmolalities correlated positively with external salinity, from 545.4+/-17.3 mOsm in 5 psu to 1185.9+/-34.6 mOsm in 41 psu. G. oceanicus hyperregulated within the 5-31.5 psu range; above 31.5 psu it hyporegulated its body fluids in comparison to the external medium. At 31.5 psu (1017 mOsm) the haemolymph concentration of G. oceanicus was isoosmotic with the habitat. The haemolymph concentrations of all the studied ions, except K(+), correlated positively with their concentrations at the various salinity.     

Non-pollen palynomorphs in the Black Sea corridor

There have been few studies of non-pollen palynomorphs (NPP) in Holocene brackish water environments. The Black Sea is one of the world’s largest and deepest bodies of stable brackish water and a natural laboratory for study of marine carbon cycling to anoxic sediments. The main NPP in the modern sediments of this brackish water sea are dinoflagellate cysts (dinocysts), acritarchs (mainly the prasinophytes Cymatiosphaera, Micrhystridium, Sigmopollis and Pseudoschizaea) and diverse fungal remains. Other NPP include colonial algae, tintinnids, copepod and cladoceran egg covers, testate amoebae and microforaminiferal linings. These NPP assemblages are similar to those in the marginal marine environment of the Pliocene St. Erth Beds (England), but have more abundant NPP, and virtually lack scolecodonts. In the Black Sea corridor, modern assemblages from areas with salinity >22‰ have higher percentages of microforaminiferal linings and fewer prasinophytes, colonial algae and fungal spores. Prasinophytes dominate only in mid-Holocene sediments, during a 2000 years interval of sea level transgression and sapropel deposition. Early Holocene sediments have lower dinocyst diversity, increased fresh–brackish water colonial algae (Pediastrum spp. and Botryococcus braunii), zygnemataceous spores and desmids (including Zygnema, Cosmarium), ostracod linings and fewer foraminiferal linings. These assemblages are similar to those in the Baltic Sea where the annual salinity is about 6–8‰.     

Noding of Cyprideis torosa valves (Ostracoda) – a proxy for salinity? New data from field observations and a long‐term microcosm experiment

Cyprideis torosa (Jones , 1850) (Ostracoda, Crustacea) is one of the most common marginal marine ostracod species in the Northern hemisphere. We investigate the relationship between variable noding of its valves and salinity as well as Ca²⁺ concentration in the ambient water, analysing populations from an in vitro experiment and field data from the southern Baltic Sea coast. There is a clear negative linear correlation between the proportion of noded individuals from our microcosms and salinity. Deficiency of Ca²⁺ causes heavier noding in laboratory cultures. The same effect can be seen in the field, however, the increase of noded individuals with falling salinity appears to be stepped, not linear. This pattern probably reflects the ability of the animals to wait some time until better salinity conditions occur within the highly variable conditions of estuaries and lagoons. At the southern Baltic Sea coast, proportions of more than 20% noded valves within a C. torosa population indicates salinities of up to 2 psu, up to 10% noded valves indicate salinities between 2 and 7 psu, and the lack of noded valves salinities > 7 psu. Stable salinity conditions as in the studied microcosms cause a shift of these salinity limits to 5 and 14 psu approximately but in a linear relationship between salinity and proportion of noded individuals. Hence, athalassic populations from more stable water bodies should be used for continuous and more detailed salinity trend reconstructions. Deficiency of Ca²⁺ (approximately < 120 mg/l) effects up to about 20% more noded individuals than in water with same salinity but with higher Ca²⁺ concentrations. The reproduction rates within the microcosms indicate a salinity optimum of C. torosa eggs of 8 psu whereas the optimum of the adults seems to be at least 14 psu (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effect of temperature and salinity on acetylcholinesterase activity, a common pollution biomarker, in Mytilus sp. from the south-western Baltic Sea

Acetylcholinesterase (AChE) activity is a well established biomarker to monitor environmental pollution caused by neurotoxic compounds, such as organophosphorus and carbamate pesticides. The presence of these compounds results in a measurable inhibition of the enzyme. This has been shown for different marine species including blue mussels. Besides pollution, environmental variables may also have a direct or indirect effect on AChE activity, particularly in estuarine and brackish water environments. To assess the impact of abiotic factors on the AChE activity the seasonal course of gill AChE activity was followed in relation to temperature and salinity in Mytilus sp. collected from the south-western Baltic Sea. In addition, the effect of salinity on AChE activity was investigated in an experimental study. The AChE activity showed significant seasonal differences with maximum activities during the summer period and minimum activities in winter. These changes correlate significantly (p<0.001) and positively with water temperature. The experimental exposure of Mytilus sp. to different salinities (5, 7, 16 or 20 psu) resulted in changes in the gill AChE activity. Empirical orthogonal function (EOF) analysis revealed that AChE activity was significantly and negatively correlated with salinity (p<0.01). These results clearly demonstrate the need to consider the potential influence of temperature and salinity on AChE activity when applying this biomarker to monitor exposure to and effect of neurotoxic substances in estuarine and brackish water blue mussels.     

Elongation and mat formation of <Emphasis Type="Italic">Chara aspera</Emphasis> under different light and salinity conditions

Former laboratory results indicate that shoot elongation at low light intensities of Chara aspera is absent already at 10 psu which is within the physiologically optimal salinity range for brackish water populations. To investigate if similar restrictions occur in the field, density and morphology of C. aspera were compared between three freshwater and three brackish water sites along its depth range. The lower depth limit of C. aspera varied considerably among sites (30–600 cm) related to turbidity. Light availability at the lower depth limit corresponded to about 15% of surface irradiance in freshwater and brackish water with lower salinity (3.4 psu). Total length increased and fresh weight:length ratio decreased with depth at these sites indicating shoot elongation related to lower light availability. Due to shoot elongation, light availability was far higher at the upper parts of the shoot than at the bottom in the turbid sites. Light availability at the lower depth limit was higher (about 40%) at two sites with higher salinity (7–8 psu), where no shoot elongation was observed at the lower depth limit. Instead, the plants were stunted and often covered with filamentous algae or shaded by other rooted submerged macrophytes indicating competitive disadvantages of C. aspera at higher salinities. As growth in high densities (mat formation) exposes the plants to severe self-shading, it is suggested that shoot elongation is a prerequisite to mat formation. Dense vegetation of C. aspera was found only in freshwater and brackish water with lower salinity. Single, richly branched plants occurred in clearwater sites with higher salinity. C. aspera was not found in “double stress” environments with both high turbidity and high salinity: We asume that the species is a poor competitor under these conditions. Our results indicate that morphological differences between freshwater and brackish water populations of C. aspera are at least partly explained by salinity rather than genetic differences.     

In situ study of relative electron transport rates in the marine macroalga <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> in the Baltic Sea at different depths and times of year

The brown alga Fucus vesiculous is one of the few marine species in the Baltic Sea. Fucus vesiculosus shows high morphological and physiological variability as a response to its environmental conditions. The salinity in the Baltic Sea is 4–5 psu, compared to 35 psu in the Atlantic. Photosynthesis of algae is usually measured after collection and transportation to constant culture conditions. However, in this study, relative photosynthetic electron transport rates, calculated from chlorophyll a fluorescence parameters were compared in algae collected from 1 and 4 m depths by SCUBA divers. Measurements of light response curves from the same individuals of F. vesiculosus at different depths and times of the year have, to our knowledge, not been made previously. Measurements were performed on four different occasions during the spring of 2005 (25 February, 3 and 29 April, and 26 May) in the Baltic Sea, using rapid light curves generated with a Diving PAM. In addition, samples were collected for photoinhibition studies in the laboratory. The light response curves obtained in situ at 1 and 4 m depths for F. vesiculosus showed lower values of light saturation with depth. When algae from 1 and 4 m depths were exposed to high irradiances of photosynthetically active radiation (1,400 μmol photons m⁻² s⁻¹), algae from 1 m depth showed a higher degree of photoinhibition in comparison to algae from 4 m depth.     

Cleavage of DNA by brown algal polyphenols

Extracts of marine algae have been tested to determine their ability to cleave DNA. The species producing positive results wereAscophyllum nodosum, Fucus serratus, F. spiralis, F. vesiculosus, Halidrys siliquosa andHimanthalia elongata. Partial purification of each extract by dialysis against water revealed that the active compounds in each species were high molecular weight polyphenols.     

A fundamental difference between macrobiota and microbial eukaryotes: protistan plankton has a species maximum in the freshwater-marine transition zone of the Baltic Sea

Remane's Artenminimum at the horohalinicum is a fundamental concept in ecology to describe and explain the distribution of organisms along salinity gradients. However, a recent metadata analysis challenged this concept for protists, proposing a species maximum in brackish waters. Due to data bias, this literature-based investigation was highly discussed. Reliable data verifying or rejecting the species minimum for protists in brackish waters were critically lacking. Here, we sampled a pronounced salinity gradient along a west–east transect in the Baltic Sea and analysed protistan plankton communities using high-throughput eDNA metabarcoding. A strong salinity barrier at the upper limit of the horohalinicum and 10 psu appeared to select for significant shifts in protistan community structures, with dinoflagellates being dominant at lower salinities, and dictyochophytes and diatoms being keyplayers at higher salinities. Also in vertical water column gradients in deeper basins (Kiel Bight, Arkona and Bornholm Basin) appeared salinity as significant environmental determinant influencing alpha- and beta-diversity patterns. Importantly, alpha-diversity indices revealed species maxima in brackish waters, that is, indeed contrasting Remane's Artenminimum concept. Statistical analyses confirmed salinity as the major driving force for protistan community structuring with high significance. This suggests that macrobiota and microbial eukaryotes follow fundamentally different rules regarding diversity patterns in the transition zone from freshwater to marine waters.     

Does scope for growth change as a result of salinity stress in the amphipod Gammarus oceanicus?

Physiological performance (feeding, metabolism, growth and excretion) across a broad range of salinity (5-30 psu) were determined for the benthic amphipod Gammarus oceanicus, a species of marine origin inhabiting brackish waters of the southern Baltic Sea. Feeding rates decreased with increasing salinity, whereas the nutritive absorption efficiency increased. Faeces production and ammonia excretion rates decreased strongly from the lowest to the highest salinity by 60% and 58%, respectively. Increasing salinity was accompanied by a reduction in the metabolic rate from 438 J g^− 1 dry wt d^− 1 (5.1 mW g^− 1) at 5 psu to 245 J g^− 1 (2.8 mW g^− 1) at 30 psu. Individuals were able to maintain a positive energy balance at all experimental salinities. The greatest values for scope for growth were recorded at the environmental salinity (7 psu) with a mean of 769 J g^− 1 dry wt d^− 1 (8.7 mW g^− 1).     

Spiniferites cruciformis: a fresh water dinoflagellate cyst?

Palynological studies of cored lacustrine sediments from the late Quaternary of Lake Kastoria, northern Greece, revealed a Late Glacial interval with abundant dinoflagellate cysts. Cyst assemblages include two identifiable species, Spiniferites cruciformis and Gonyaulax apiculata. The presence of the fresh water species G. apiculata is consistent with the lacustrine setting of these deposits, but that of S. cruciformis is anomalous. Previously, this species has only been recorded in abundance from presumed brackish marine sediments from the Black Sea and Marmara Sea sediments where geochemical data clearly record brackish salinities. Therefore, it has been regarded as a low salinity cyst type with a wide range of morphological variation that some workers have suggested to reflect salinity fluctuations. Specimens from Greece display only part of the range of morphological variability previously described from these (brackish) marine settings. Encountered morphological variation includes ellipsoidal/pentameral and cruciform endocyst shapes with rare intermediate shapes, and highly variable septa development. Specimens characterized by extremely reduced ornamentation known from (brackish) marine environments have not been recorded. Our records of S. cruciformis indicate that: (1) it could thrive in fresh water conditions; and (2) that apparently most of the strong morphological variations of the cysts are an intrinsic phenomenon for this taxon, and may only partly be linked to salinity variations as suggested earlier. We suggest that S. cruciformis essentially is a fresh water taxon, and that its records in (brackish) marine environments, with the exception of specimens with strongly reduced ornamentation, may be due to transportation, to short-lived fresh water surface conditions in such environments, or to tolerance of the species to brackish conditions.     

Influence of changes in salinity and light intensity on growth of phytoplankton communities from the Schelde river and estuary (Belgium/The Netherlands)

In the Schelde continuum, a succession in the phytoplankton community is observed along the transition from the river to the freshwater tidal reaches of the estuary and from the freshwater to brackish reaches of the estuary. The goal of this study was to experimentally evaluate the contribution of changes in salinity and light climate to this succession. In summer 2000 and in spring 2001, phytoplankton communities from the river, the freshwater tidal reaches and the brackish reaches of the estuary were incubated under high or low light intensities and exposed to a change in salinity. HPLC analysis was used to evaluate the response of different algal groups to changes in light intensity and salinity. When incubated at a light intensity corresponding to the mean underwater light intensity of the freshwater tidal reaches, growth of phytoplankton from the river as well as from freshwater tidal reaches was significantly lower than when incubated at a light intensity corresponding to the mean underwater light intensity of the river. The phytoplankton community from the freshwater tidal reaches did not appear to be better adapted to low light intensities than the phytoplankton community from the river. Although diatoms were expected to be less sensitive to a reduction in light intensity than green algae, the opposite response was observed. Freshwater and brackish water phytoplankton were negatively affected by respectively an increase or decrease in salinity. However, the effect of salinity was not strong enough to explain the disappearance of freshwater and brackish water phytoplankton between a salinity of 0.5 and 10 psu, suggesting that other factors also play a role. In the freshwater phytoplankton communities from the river and the freshwater tidal reaches, green algae and diatoms responded in a similar way to an increase in salinity. In the brackish water phytoplankton community, fucoxanthin displayed a different response to salinity than lutein and chlorophyll a.     

The growth of pikeperch (Sander lucioperca L.) and perch (Perca fluviatilis L.) under different water temperature and salinity conditions in the Curonian Lagoon and Lithuanian coastal waters of the Baltic Sea

Pikeperch and perch perform seasonal migrations between the Curonian Lagoon and the coastal waters of the Baltic Sea. The Curonian Lagoon is a freshwater basin, while salinity in the coastal waters varies between of 4.9–6.8 psu. In the Curonian Lagoon water temperature is generally higher than in the coastal waters. Field studies of growth and condition characteristics of pikeperch and perch were carried out in these water bodies with the aim to estimate growth differences of the two fish species under different salinity and temperature conditions. Additionally, an experimental study of the impact of salinity on the growth of perch young-of-the-year (YOY) was performed to test the hypothesis that a brackish environment positively influences percids. Field observations revealed that body length, condition factor, fatness coefficient and fat content in muscles were significantly higher in individuals inhabiting the cooler, brackish waters of the Baltic Sea than in individuals inhabiting the Curonian Lagoon. A positive effect of low salinity on growth was also established in the experimental study. Hence, the study results suggest that under certain temperature conditions, brackish waters beneficially affect the growth of pikeperch and perch.