The reproductive biology of the dab Limanda limanda (L.) in the North Sea: Seasonal changes in the ovary

Oogenesis in the dab is described in six easily identifiable stages: oogonium (Stage I), primary oocyte (II), primary vitellogenesis (III), secondary vitellogenesis (IV), tertiary vitellogenesis (V) and hyaline oocyte (VI). The annual reproductive cycle in female dab may be divided into four morphologically and histologically distinct periods: prespawning (October–January), spawning (February–April), postspawning (May–June) and resting (July–September) period.     

The reproductive biology of the dab Limanda limanda (L.) in the North Sea: gonosomatic index, hepatosomatic index and condition factor

Measurements have been made of the annual cyclical changes in gonosomatic index (GSI), hepatosomatic index (HSI) and condition factor ( K ) in male and female dab Limanda limanda (L.). There is a direct correlation between HSI and K , and an inverse correlation of these factors to GSI but with a three month lag. It was also found that the lower ('ventral') ovaries were invariably heavier than the upper ('dorsal') ovaries at all times, and a similar relationship was found in testes prior to and during the spawning season.     

The karyotype of the dab (Limanda limanda L.)

In the dab ( Limanda limanda L.), the diploid number is 46 and the karyotype consists of 23 pairs of chromosomes, all of which are telocentric or subtelocentric, so that the total number of chromosome arms is 46 (AN-46).     

Seasonal changes in the common dab, Limanda limanda (L.) in Isle of Man Waters

Otolith accretion was greatest in summer when opaque bands were laid down and slowest in winter when hyaline rings were formed. Food intake was minimal in winter and maximal in summer. Liver weight was high in summer and winter and low in spring after spawning. Growth in length was higher in summer than winter. Gonad weight was greatest in winter for both sexes. Spawning occurred from February to April. After spawning males recovered earlier than females and testes ripened earlier than the ovaries. From winter to summer, the sea temperature increased from 7.5–13.0°C; the day length from 7–17h; and the salinity varied from 34.07−34.36%.     

Seasonal changes in frequency of diseases in dab,Limanda limanda,from the southern North Sea

Infestation rates of flatfishLimanda limanda from the southern North Sea (German Bight) were investigated in 1981 and 1982. At three stations, including a dumping area of acid-iron wastes northwest of Helgoland, dabs were examined for the occurrence of lymphocystis and epidermal papillomas. Marked seasonal changes of infestation rates were observed with maxima of diseased fish in March and May (epidermal papillomas 2 % in 1981, 4.1 % in 1982; lymphocystis 7.5 % in 1981, 17.2 % in 1982). Infestation minima were found from June to October. Differences in incidence rates were observed between stations in winter and spring, but not in summer. Both diseases turned out to be size-dependent. Seasonality of the two diseases is discussed in relation to spawning cycle and water quality, including pollution.     

Some observations on the seasonal distribution of dab,Limanda limanda, in the southern North Sea

The seasonal changes in the population structure of dab were observed in intervals of four to six weeks from January 1993 to November 1994 at a fixed station northwest of Helgoland. The sex distribution was characterized by a reduced number of males during autumn and winter. In the autumn of both years, an immigration of large females was evident. During long periods in spring and summer, the population structure remained relatively constant. The observations indicate that dab in the Helgoland region carry out a seasonal migration between coastal waters and the sampling area. Seasonal changes of the population structure occurred at a relatively slow rate.     

Assessment of some innate immune responses in dab (Limanda limanda L.) from the North Sea as part of an integrated biological effects monitoring

The marine flatfish dab (Limanda limanda), which lives in direct contact with contaminated sediments, is frequently used as a sentinel species in international monitoring programmes on the biological effects of contaminants. In this study, immune responses were recorded as indicators of sublethal chronic effects of contaminants, in addition to measurement of the induction of mono-oxygenase ethoxyresorufin O-deethylase (EROD) in liver cells, the inhibition of acetylcholin esterase (AChE) in muscle and a quantification of grossly visible diseases and parasites. In total, 336 dab were analysed from five sampling areas in the North Sea, including the German Bight, the Dogger Bank, the Firth of Forth, and two locations close to oil and gas platforms (Ekofisk and Danfield). When considering plasma lysozyme levels, pinocytosis and respiratory burst activity of head kidney leucocytes, a clear gradient could be observed with decreased levels in individuals collected from the Firth of Forth and locations near the oil or gas platforms compared with dab from the Dogger Bank or the German Bight. Individuals with induced EROD activity displayed reduced lysozyme and respiratory burst activities. Lysozyme levels were also reduced in dab with lymphocystis or with nematodes. The data obtained indicate that the assessment of innate immune parameters in a monitoring programme provides supplementary information about immunomodulatory effects associated with the exposure of fish to contaminants. In particular, concentrations of plasma lysozyme, which can be analysed in an easy and inexpensive assay, are considered to be an appropriate parameter for use in a battery of other bioindicators. Communicated by H. von Westernhagen and A. Diamant     

Development of 30 microsatellite markers for dab (Limanda limanda L.): a key UK marine biomonitoring species

Dab (Limanda limanda) are the principal target fish species in offshore biomonitoring programmes in the UK; however, detailed knowledge of genetic structure and connectivity among sampling locations is unavailable. Here, the isolation and characterization of 30 polymorphic microsatellite loci for dab is described. The number of alleles per locus ranged from 2 to 42, with observed heterozygosities ranging from 0.089 to 1. These loci will enable high resolution of genetic population structure and dynamics of dab around the British Isles.     

The feeding strategy of dab Limanda limanda in the southern North Sea: linking stomach contents to prey availability in the environment

The feeding strategy of dab Limanda limanda was investigated within a predefined research area in the southern North Sea for 2 years. The aims were: 1) to establish the relationship between prey availability and dab abundance, distribution, body condition and prey selection, and 2) to determine the underlying functional processes of prey choice in dab. Temporal and spatial trends of prey availability in relation to dab abundances, distribution, condition and prey selection were analysed. Temporal trends in dab condition, percentage of empty stomachs and numbers of ingested prey were apparent. In particular in the winter period 2000–2001 dab were in poor condition and prey items were only rarely found in stomachs. Similarly prey availability decreased in the environment while the abundance of dab increased. Thus, temporal changes in the nutritive state were partly thought to be attributed to density‐dependent effects. The stomach composition of dab during the two consecutive summers was mainly dominated by Pariambus typicus and Ophiura albida while in the winter period only O. albida made a significant contribution to the stomach content. Analysis of prey characteristics indicated that only prey densities in the environment significantly influenced prey choice in dab, while other characteristics such as the position of the prey in the habitat, its palatability or mobility did not have significant effects. The feeding strategy of dab thus was highly opportunistic, although trends were apparent that suggested that buried living fauna was less likely to be ingested, which was also observed in previous studies.     

Maximum daily ration and the pattern of food consumption in haddock, Melanogrammus aeglefinus (L.), and dab, Limanda limanda (L.)

Food intake was measured under conditions of continuous food availability for haddock, Melanogrammus aeglefinus (L.), and dab, Limanda limanda (L.). Haddock consumed approximately 75% of all food during the day, with a peak of feeding at dusk and an inferred peak at dawn; peaks of feeding were associated with larger meals. By contrast, dabs were mainly night-time feeders with daily consumption rates that were approximately 40% lower than those shown by haddock. Comparison of haddock maximum daily ration with published estimates of consumption in the wild indicate that wild haddock feed at between 15 and 22% of their maximum capacity. Using published data on gastric evacuation, the relationship between meal size and stomach residuum was investigated. There was no detectable relationship between these two variables.     

Emptying rates of single and double meals of different food quality from the stomach of the dab, Limanda limanda (L.)

In previous studies on gastric emptying time in Limanda , data were obtained which predict that food will empty from the stomach according to: after a temperature sensitive delay. This curve gives an excellent prediction of the emptying of both artificial and natural food items given as single meals when stomachs are sampled directly. However, when two meals are given 3 h apart, emptying rate depends on whether the two meals remain separate or are allowed to mix by omitting the binding agent. In the absence of a binder, both meals are slowed so that the overall emptying rate is as predicted by the equation. When binder is present, the first meal is not delayed and the overall gastric emptying rate is increased 35%.     

Gastric emptying in Limanda limanda (L.) and the return of appetite

Limanda limanda (35–225 g) fed during daytime when trained to use demand feeders under natural photoperiods. Under continuous illumination different feeding rhythms developed which were detected by periodogram analysis. Interfeeding periods (P hours) varied with body weight (W grams) and temperature (T° C) over the range 6.5–15° C as: In P=4.4 – 0.167 (In W) – 0.05 T. Daily food intake (D grams increased with both temperature and size (In D = 0.099 T + 0.579 In W – 3.49). Relative daily intake (R as % body weight, b.w) increased with temperature but decreased with size (In R = 0.099T–0.41 In W + 1.05). These changes in daily intake are primarily caused by changes in feeding frequency; the amount eaten at a meal is relatively constant save at low temperatures (6.5° C) when appetite is suppressed. Dilution of the food with kaolin (from 4.8 to 2.2 kcal/g) did not cause a compensatory increase in meal size or frequency. Gastric emptying time (GET hours) measured by X-radiography changed with body weight, temperature and relative meal size (M % b.w.) as In GET = 0.68 In M + 0.39 In W – 0.035T + 1.46. With the exception of the lowest temperature (6.5° C) interfeeding periods closely followed GET (P = 1.09 GET – 2.6) supporting the proposal that stomach fullness is a major factor controlling appetite in the dab.     

Regional patterns in prevalence of principal external diseases of dab Limanda limanda in the North Sea and adjacent areas 1992-1997

The prevalence and spatial distribution of major diseases of dab Limanda limanda in the North Sea and adjacent areas were studied in the summers 1992 to 1997. Areas covered were the North Sea, Irish Sea, northern and northeastern British Waters and the English Channel. The diseases studied were lymphocystis, epidermal hyperplasia/papilloma and skin ulceration. To standardise data, results were analysed for females >15 cm (>3 yr old). Data were subjected to median polish, and additive, extended and additive plus multiplicative models were applied to best account for effects of region and year. Annual differences in disease prevalence were low whilst differences between areas were pronounced. For lymphocystis higher prevalence was observed in the northwestern sector of the North Sea, at the northern tip of Scotland and in an area south of Iceland. Prevalence was low in the Irish Sea, the English Channel and the southern North Sea, and intermediate in the German Bight. For epidermal hyperplasia/papilloma, levels were low at Icelandic stations, in the northern Irish Sea, in the southern North Sea and the English Channel, whilst levels were high in the northwestern part of the North Sea and the German Bight. Elevated levels of skin ulceration were found on the Dogger, at 1 station in the Irish Sea (off Sellafield) and at 1 station to the south of Iceland. Lower levels were detected west of Iceland. Prevalence in all other areas was intermediate. It is concluded that a detailed analysis of available data on disease prevalence and putative causative factors is desirable and, given the good availability of data, would be a promising step forward toward elucidating possible cause and effect relationships between diseases and anthropogenic factors.     

Small-scale variability in growth and condition of dab Limanda limanda (L.) larvae in relation to an Irish Sea tidal-mixing front off the west coast of the Isle of Man

The small-scale variability in condition and growth of dab Limanda limanda larvae was investigated in relation to a tidal-mixing front (Manx West Coast front) off the west coast of the Isle of Man, Irish Sea. RNA:DNA ratios and otolith microincrement analysis were used to estimate nutritional status and recent growth rates of larvae captured on 5 May 1999 in mixed, frontal and stratified waters associated with this front. RNA:DNA ratios were significantly higher at the Manx West Coast front compared to mixed waters, although recent growth rates did not vary significantly with water column stability. More favourable feeding conditions associated with frontal waters from previously reported large-scale investigations in the Irish Sea were reflected on a smaller spatial scale. Although frontal accumulation of dab larvae was not observed in the current study, dab larvae found in the stratified and frontal waters were significantly larger than those captured in the mixed waters. This pattern could suggest an offshore movement of dab larvae into more favourable waters as they develop.     

Ethoxyresorufin O-deethylase (EROD) activity in the liver of dab (Limanda limanda L.) and flounder (Platichthys flesus L.) from the German Bight. EROD expression and tissue contamination

Ethoxyresorufin O-deethylase (EROD) activity was measured in the liver of dab (Limanda limanda) and flounder (Platichthys flesus) from the German Bight (southern North Sea) and compared with muscle and liver polychlorinated biphenyl (PCB) concentrations in an attempt to relate EROD activity to PCB body burden. In none of the different datasets (species-, tissue- or matrix-dependent) was a significant (P<0.05) correlation between PCB tissue contamination and EROD activity found. Yet EROD activity was significantly correlated with polycyclic aromatic hydrocarbons (PAH) levels (phenanthrene, fluoranthene, pyrene) in muscle tissue, indicating a possible dependence of EROD expression on other ubiquitous organic contaminants, thus making it a suitable biomarker for general pollution. Received: 14 April 1999 / Received in revised form: 10 July 1999 / Accepted: 15 July 1999     

The reproductive biology of the horse mackerel Trachurus trachurs (L.) in the North Sea and English Channel

In the horse mackerel the gametes develop asynchronously and are released in batches. The maturity cycle is characterized by a rapid post-spawning recovery period and a relatively long period of oocyte maturation. Mean gonad weight reached a peak in June and intraovarian hyaline oocytes occurred from May to August. Determination of fecundity was complicated by resorption of oocytes both before and after spawning, and by the fact that resting and developing oocytes were not clearly separated by size. An estimate of'potential fecundity'was obtained by a combination of histology and oocyte measurement. Plankton surveys showed that a major spawning area exists in the Southern Bight of the North Sea, with eggs occurring from May to late August. An estimate of annual egg production was combined with the fecundity data to give an estimate of stock size of about 500 000 tons.     

Some effects of temperature, meal size and body weight on gastric evacuation time in the dab Limanda limanda (L)

Gastric emptying time in the dab, Limanda limanda , has been studied using an X-ray technique. The addition of 25 % barium sulphate to a test meal did not significantly affect the transit time. Lowering the experimental temperature from 16.4 to 8.5°C markedly decreased gastric evacuation time. An increase in ration size led to an increase in the time required to empty the stomach and also to increase the amount of food digested per unit of time. For any given ration size, expressed as per cent body weight, the larger the animal the longer is the time required for evacuation of that meal. We suggest that the food intake per day, as a percentage of live body weight, will be smaller for larger L. limanda in the wild.