Sub-lethal exposure to ultraviolet radiation reduces prey consumption by Atlantic cod larvae (Gadus morhua)

Levels of ultraviolet-B radiation (UV-B: 280–315 nm) reaching the earth’s surface have increased over the past few decades due to ozone depletion. It is well documented that exposure to UV-B radiation increases mortality in marine fish larvae. However, few studies have examined sub-lethal effects of UV-B radiation such as, for example, the possibility that it affects prey consumption by fish larvae. Atlantic cod larvae were exposed to a sub-lethal level of ultraviolet-B radiation (UV-B: 280–315 nm) for 15 h. After the exposure, rotifers (4/mL) were fed to cod larvae at three different post-exposure intervals (20 min, 3 and 6 h). Trials were replicated three times for each post-exposure interval. The number of rotifers in the gut and the percent of empty guts (number of fish with empty gut/number of fish examined × 100) were analyzed. Results were compared to those of unexposed fish (the control treatment). UV-B exposed cod larvae had consumed significantly fewer rotifers than control fish at all post-exposure intervals. There was no significant difference in the frequency of empty guts between fish in the UV-B treatment versus fish in the control treatment at any of the post-exposure time points (although the difference at 20 min post-exposure was borderline significant). These observations suggest that cod larvae exposed to sub-lethal levels of UV-B have lower net energy gain which may lead to reduced growth rate and possibly poorer survival.     

Oxidative stress and changes in locomotor behavior and gill morphology of Gambusia affinis exposed to chromium

Sublethal effects of chromium trioxide on mosquito fish, Gambusia affinis, were carried out for 28 days on day 4, 8, 12, 16, 20, 24, and 28. The sublethal effects of chromium (LC₁₀, 77.62?mg?L^?1) on antioxidant enzymes, thiobarbituric acid reactive substance (TBARS), locomotor behavior, and gill morphology were studied. The antioxidant enzymes like superoxide dismutase (SOD) and catalase were induced throughout the exposure span of 28 days. Percent induction was also increase with the days of exposure. Lipid peroxidation product, malondialdehyde was enhanced in the viscera tissue of chromium-exposed fish. Perturbation of locomotor behavior like decrease in distance travelled (m?min^?1) and in swimming speed (cm?s^?1) was observed in exposed fish. Chromium toxicity caused alterations in gill morphology like hypertrophy and hyperplasia in secondary lamellae followed by detached epithelium with severe necrosis. It can be concluded from the results that Cr intoxication in Gambusia fish, caused induction in antioxidant enzymes and lipid peroxidation, plus changes in gill morphology and locomotor behavior.     

Effects of UV radiation of different wavebands on pigmentation, 15N-ammonium uptake, amino acid pools and adenylate contents of marine diatoms

Three marine diatoms Lauderia annulata Cleve, Odontella sinensis (Greville) Grunow and Thalassiosira rotula Meunier were exposed to ultraviolet (UV) radiation of different wavebands under controlled laboratory conditions (0.035 vol% CO₂, 18 °C). Several changes in the patterns of pigments in these organisms were seen depending on the waveband of UV radiation and species examined. UV-B and UV-B plus UV-A radiation led to a reduction in the overall pigment content of all three diatoms. The uptake of ¹⁵N-ammonium was less affected by 5-h UV-A (WG 320) but significantly reduced after UV-B and UV-B plus UV-A exposure. The pattern of free amino acid pools varied depending on the applied UV wavebands and the tested diatom. The main protein-bound amino acids of T. rotula decreased after 5-h UV irradiance except leucine. Contents of adenosine 5′-mono-, di-, and triphosphate (AMP, ADP and ATP) were affected differently by UV radiation; ATP values increased at the end of UV-B and UV-B plus UV-A exposure. These results have been discussed with reference to the impact of the different UV sources and the influence on the nitrogen metabolism in connection to pigments and supply with energy. Received: 13 May 1997 / Accepted: 11 October 1997     

Effects of ultraviolet radiation on skin structure and ultrastructure in Caspian Sea Salmon,Salmo trutta caspius,during alevin stage

Ozone reduction enhances the amount of ultraviolet (UV) radiation reaching the aquatic ecosystems, which can induce adverse effects on fish. In order to study the UV radiation effects on skin structure and ultrastructure, the alevins of Salmo trutta caspius were exposed to two different doses of UV radiation (low doses of: 60 µW cm^?2 UVC; 100 µW cm^?2 UVB; and 40 µW cm^?2 UVA and high doses of: 90 µW cm^?2 UVC; 130 µW cm^?2 UVB; and 50 µW cm^?2 UVA) for 15 min once a day in dark condition. Alevins with no exposure to either solar light or UV exposure served as the control group. No mortality was observed in the control group, whereas all of the alevins died in the 9th day in both UV exposure groups. Light microscopic and scanning electron microscopic studies showed significant reduction in the epidermis thickness, number of skin mucous cells, and percentage of mucus secreting cells in both UV exposure groups. Sunburn cells, lifting epidermis from basal membrane, and disappearance of pavement cell microridges were also observed. Our results showed that UV radiation caused intensive skin cell lesions in the Caspian salmon alevins. Since the skin cells of trout alevins possesses essential roles in respiration, osmoregulation, excretion and defense, the observed damages to the skin could be attributed to their sudden mortality in the UV exposure.     

Laser Radiation Can Retard Leaf Senescence of Isatis Indigotica Seedlings Exposed to Elevated UV-B

以菘蓝(Isatis indigotica Fort)幼苗为实验材料,比较研究了He-Ne激光对增强UV-B(辐射强度:10.08kJ·m-2;照射时间8h·d-1;共照射8d)辐射损伤菘蓝叶片衰老的延缓作用.结果表明:与对照相比,UV-B辐射可导致环丁烷嘧啶二聚体(CPD)形成,显著降低蛋白酶、转氨酶、游离氨基酸、可溶性蛋白质含量,同时降低了叶绿素含量、气孔导度、水分利用率和生物量;而菘蓝幼苗经UV-B辐射处理后再用激光辐照(波长:633nm;功率密度:5.23mW·mm-2;束径:1.5mm;照射时间:5min·d-1),叶片中CPD含量显著降低,蛋白酶、转氨酶、游离氨基酸、可溶性蛋白质含量、叶绿素含量、气孔导度、水分利用率和生物量与单独UV-B处理组相比显著提高.上述结果表明激光辐照可以显著延缓UV-B损伤菘蓝叶片的衰老。     

Factors influencing the distribution and coexistence of Pachygrapsus crassipes and Hemigrapsus oregonensis (Decapoda: Grapsidae) in a California salt marsh

It has recently been demonstrated that acute midultraviolet irradiation (UV-B, 290 to 320 nm) of the marine copepod Acartia clausii results in reduced survival and fecundity. In the present study, immature late copepodites were separted by sex and exposed to three UV-B exposure levels (0, 25, and 50 effective_DNA Jm^-2). The irradiated copepods were then reared to sexual maturity, and adult virgin survivors were mated according to prior exposure, supplying 7 different types of mating crosses (M0F0, M0F25, M25F0, M25F25, M0F50, M50F0, M50F50). Six replications of each cross were maintained for the duration of an experiment. Two experiments are documented. There was a significant effect of UV-B radiation on the survival of the parents as well as a reduction in the number of eggs and the number of living nauplii produced. For each experiment, 30 nauplii from each type of cross were separated, reared to maturity and counted, providing information on the survival capability of non-irradiated offspring from the 7 different mating types. Previous exposure of parental stock did not significantly affect the survival, and development to maturity, of these nauplii 15 d after separation.     

Histology of Sculpin spp. in East Greenland. II. Histopathology and trace element concentrations

Abstract

For many years, the sculpin has been utilized as a sentinel monitoring species for anthropogenic impacts on the marine environment. To further develop its potential as a screening model body burden of several trace elements, including cadmium (Cd), copper (Cu), mercury (Hg), lead (Pb) and selenium (Se), were investigated. In addition, histopathology in shorthorn sculpins (Myoxocephalus scorpius; n?=?20) and fourhorn sculpins (Myoxocephalus quadricornis; n?=?10) were examined at three sites around the Scoresby Sound settlement in Central East Greenland. Fourhorn sculpins caught at the most distant site from the settlement, contained significantly higher hepatic Cu and Cd concentrations as well as significantly elevated gill Pb levels compared to the shorthorn sculpins collected at two other sites closer to the settlement. Histological examinations showed that fish with significantly higher concentrations of hepatic Cd and Se exhibited greater nuclear alterations, interstitial hyperplasia/hypertrophy, interstitial mononuclear infiltrations and granulomas. Further, fish with higher Cd and Cu gill tissue levels displayed a significantly higher number of cytoplasmic alterations and lamellar epithelium lifting, hypertrophic and hyperplastic epithelium along with mucus cell hyperplasia. While the presence of liver lesions were not species or sex-specific, the presence of gill lesions decreased in the order female fourhorn sculpins?>?female shorthorn sculpins?>?male fourhorn sculpins?>?male shorthorn sculpins. Hepatic Hg concentrations exceeded known lowest observable effect doses (LOED) for fish (0.1–0.5?μg g^?1 ww) in 27% of fish, while liver Cd residues in 80% exceeded LOED (0.42–1.8?μg g^?1 ww). Based upon these results, data suggest that using the sculpin as a valuable sentinel fish species histopathology may serve as a reliable tool for assessing marine ecosystem exposure to trace metals. However, confounding physiological and ecological factors also need to be considered.     

Biomarkers as tools to assess the chronic toxicity of ammonia in the juvenile Mugil cephalus

With juvenile fish as the subject, the effects of low concentration ammonia on antioxidant system were studied using Mugil cephalus. Samples of gill and liver tissue were obtained from 0.35, 0.70, 1.5 and 3?mg/L ammonia groups at 0, 5, 10, 15 and 20 days of exposure, at which times the biomarkers were measured. Results showed that gill malondialdehyde (MDA) content exhibited an initial significant increase (p?≤?0.05) at unionised ammonia concentrations of 0.70, 1.5 and 3.0?mg/L on day 5, followed by subsequent declines, while liver MDA levels exhibited significant increases (p?≤?0.05) at unionised ammonia concentrations of 1.5?mg/L starting on day 10 and at 3.0?mg/L starting on day 5. With exposure to ammonia at different concentrations, Na⁺-K⁺-ATPase activity in liver and gill decreased over time. The Na⁺-K⁺-ATPase activity was negatively related to ammonia concentration from 0.70 to 3.0?mg/L. Overall, our results show that MDA and Na⁺-K⁺-ATPase, evaluated here as potential biomarkers of ammonia exposure, exhibited responses to sublethal concentrations of ammonia that were concentration dependent.     

Consequences of Embryonic UV-B Exposure for Embryos and Tadpoles of the Plains Leopard Frog

Abstract: In organisms with complex life cycles, understanding the effects of environmental change requires investigating the possibility that the effect of such change on one stage will have consequences for other stages. We investigated the effects of embryonic exposure to UV-B radiation on embryo and tadpole development of the plains leopard frog (   Rana blairi ). Embryos were exposed to sunlight filtered to provide two levels of UV-B exposure (e.g., at 310 nm: high exposure is approximately 84% transmittance and low exposure is approximately 58% transmittance). Hatching success of embryos exposed to different UV-B levels did not differ. Tadpoles resulting from exposed embryos were raised at three densities. Growth and development were slower in tadpoles exposed to higher UV-B levels as embryos. Density negatively affected growth and development. Survivorship did not differ among UV-B or density treatments. Our results suggest that even if lethal effects are not exhibited at one stage, sublethal effects may manifest themselves at other stages, possibly affecting the long-term success of exposed individuals and populations.     

Effect of surfactants on phosphatase level of fresh water fish Labeo rohita

Alterations in the activity of enzymes Acid phosphatase (E.C.3.1.3.2) and Alkaline phosphatase (EC 3.1.3.1) in organs such as liver, gills and muscle of rohu following its exposure to surfactants viz. CTAB, SDS and Triton X-100 were analyzed. Different levels of exposure were given depending on the LC50 value of the surfactant used. Also, the influence of age and weight of the organisms was tested simultaneously. The activity of ACP in the tissues of fish exposed to all the three surfactants showed marked enhancement after exposure; the effect being highest in the liver followed by gill and muscle. Activity levels of ALP in different tissues of the fish exposed to the surfactants also showed an increase. Maximum increase was found in the liver followed by muscle, and gill. The increase in the levels of these enzymes indicates a stressful condition of the fish.     

Effects of cadmium on metallothionein and histology in gills of tilapia (Oreochromis niloticus) at different salinities

The objectives of this study were to evaluate the effects of sublethal cadmium concentrations on the levels of cadmium, metallothionein (MT) and histological changes in gills of East Java strain tilapia (Oreochromis niloticus) at different salinity levels. The levels of cadmium in control gills were not significantly different at 0, 5 and 10 practical salinity unit (PSU). The cadmium concentrations in gills of cadmium-exposed fish were significantly higher at 0 PSU than at 5, 10 and 15 PSU. The MT concentrations in control gills were not significantly different at 0, 5, and 10 PSU. The MT concentrations of cadmium-exposed fish were significantly higher than those in respective control groups at 0, 5 and 10 PSU. Significant gill damage occurred in fish exposed to cadmium at lower salinity. The epithelial lifting was noted at gills of fish exposed to 2.5 mg/L of cadmium at 0 PSU, and telangiectasis was observed at gills exposed to 5 mg/L of cadmium at 0 PSU. The level of gill damage decreased with increasing salinity of media. The increased MT and histological changes in gills of our findings could be a protective response of animals to toxic effect of cadmium.     

Gas–liquid chromatography for fenvalerate residue analysis: alterations in the ionic concentration and associated adinosine triphosphatases in fish,Cirrhinus mrigala (hamilton)

In the present study, an attempt has been made to quantify the fenvalerate accumulated in different tissues (gill, muscle and liver) and observe changes involved in the levels of sodium, potassium and calcium ions and Na⁺–K⁺, Mg²⁺ and Ca²⁺ adenosine triphosphatase (ATPase) activities in the freshwater fish, Cirrhinus mrigala on short-term and long-term exposure to the median lethal and sublethal concentration of fenvalerate. Residue analysis using gas–liquid chromatography (GLC) technique revealed that fenvalerate accumulated in highest quantity in gill followed by liver and muscle under median lethal concentration (6?µg?L^?1). Whereas in sublethal concentration (0.6?µg?L^?1), muscle accumulated highest quantity followed by gill and liver, which might be due to the fact that fenvalerate is highly lyphophilic. The ion concentration and ATPase activity were found effected in fish exposed to lethal and sublethal concentrations of fenvalerate. Concentration of Na⁺, K⁺ and Ca²⁺ ions decreased in gill, muscle and liver on being exposed to median lethal concentration to a significant level. Whereas the changes were not highly pronounced at sub lethal level indicating low concentration of fenvalerate and its non-toxic effect at chronic exposure. Na⁺–K⁺, Mg²⁺ and Ca²⁺ ATPases activity were also found decreased in correspondence to the ionic change under median lethal and sub lethal concentrations in target tissues. This might have lead to behavioural changes and create wide-spread disturbance in the normal physiology, ultimately causing the death of the fish. The results suggest that in biomonitoring programmes, ions and associated ATPases can be a good diagnostic tool for fenvalerate toxicity.     

Studies of some biochemical changes in the tissues of Catla catla (Hamilton), Labeo rohita (Hamilton) and Cirrhinus mrigala (Hamilton) exposed to NH3-N,NO2-N and NO3-N

Biochemical changes, total proteins, glycogen, aspartate and alanine (AAT and ALAT) amino transferases were studied with exposure of sublethal concentrations of NH3-N, NO2-N and NO3-N to the freshwater fish Catla catla (Hamilton), Labeo rohita (Hamilton) and Cirrhinus mrigala (Hamilton). Depletion in the food reserves and enzyme activity was observed in all the three fish species exposed to these toxicants. Hence, the concentrations of NH3, NO2 and NO3 in water need to be monitored in water quality in aquaculture practices.     

Municipal wastewater-induced histopathological lesions in gills of freshwater fish Labeo rohita – A SEM study

The fingerlings of fish Labeo rohita were subjected to chronic (15, 30, and 60 days) and recovery (60 days) experiments for histopathological studies using sublethal concentrations (35.4%, 26.6%, and 17.7% v/v) of municipal wastewater of Tung Dhab drain, Amritsar, India. Control fish were maintained for same exposure periods using tap water. Scanning electron microscopic studies exhibited dose- and duration-dependent alterations on gill surface ultra-structure. Control fish exhibited functionally normal gills with normal mean degree of tissue change. Fifteen different types of lesions were identified out of which 9, 5, and 1 were of the stage I, II, and III, respectively. The mean degree of change values in the treated groups ranged from 14.8 ± 0.5 (17.7%, 15 days) to 116. ± 17 (35.4%, 60 days) demonstrated gill damages of slight to irreparable lesions. The recorded alterations were epithelial lifting and intraepithelial edema, reduction of water spaces, lamellar fusion, lamellar telangiectasia, swelling and fusion of filaments, sloughing of lamellar epithelium, hypertrophy and hyperplasia of mucous and chloride cells, and necrotic lamellae. The recovery experiments showed positive improvement as the intensity of damage was less than recorded for 60 days of treatment. This study clearly indicated toxicity of drain water to fish at cellular and organ level. The study recommends environmental agencies to make and enforce decisions regarding improvement of water quality of Tung Dhab drain, and further incorporating histopathological studies as an analytical tool.     

Impact of diesel oil effluent in the mucosal surface of the alimentary canal of Oreochromis nilotica (Linnaeus): a scanning electron microscopic study

Adverse effects of diesel oil on microanatomical structure of the alimentary canal of O. nilotica were studied using SEM observations. The study revealed irregular arrangement of the stratified epithelial cells along with fragmentation of the normal concentric pattem of microridges of the same cells in buccopharynx and oesophagus. The excessive secretion of mucus of buccopharynx and oesophagus were the salient changes caused by diesel oil pollution. The destruction and degeneration of the mucosal folds of stomach and intestine along with their epithelial cells exhibited a concrete hyperactivity resulting in abundant' secretion of mucus over the microridges of the epithelial cells. In the intestinal region the columnar epithelial cells showed tumefaction and microvilli of the plasma membrane of epithelial cells get heavily damaged. Disarray of the microridges of epithelial cells, excessive secretions of mucus formation of even cell sheet were the most conspicuous changes in rectum. It was concluded that chronic exposure of diesel oil may hamper the absorption of the nutrients through alimentary canal resulting into ill-growth and production of the fish.     

Survival and sex ratios of the intertidal copepod,Tigriopus californicus,following ultraviolet-B (290–320 nm) radiation exposure

The marine harpacticoid copepod, Tigriopus californicus (Baker), is a successful colonizer of supralittoral splash pools from Torch Bay, Alaska, to Baja California, Mexico. As these pools are subject to abundant amounts of direct solar radiation, it was of interest to determine the sensitivity of T. californicus to ultraviolet-B (UV-B, 290 to 320 nm) radiation. During 1980, copepods were raised under diurnal conditions in the laboratory and fed a mixture of unicellular algae and bacteria. Larval and adult stages were irradiated for 0 to 26 h on a rotating turntable under enhanced ultraviolet radiation. Following irradiation, the copepods were maintained in the culture area and checked daily for survival. All life stages of T. californicus tolerated enhanced UV-B radiation irrespective of exposure period; in contrast, survival of an irradiated planktonic copepod, Acartia clausii (Giesbrecht), was significantly lower. Additionally, there appeared to be asex ratio shift in adults that developed from the youngest naupliar stages (N₁/N₂) of T. californicus that were irradiated. Extraction and tentative identification of the pigment(s) responsible for the bright orange color of the carapace of T. californicus were performed. These pigments may function to absorb UV radiation and protect internal structures from photochemically induced damage.     

Effect of phenol on haematological components of Indian major carps Catla catla, Labeo rohita and Cirrhinus mrigala

The effect of phenol on haematological components of Indian major carps, Catla catla, Labeo rohita and Cirrhinus mrigala were observed. After exposure to sublethal concentrations of 5.17 mg l(-1), 6.06 mg l(-1) and 6.99 mg l(-1), the number of red blood cells, haemoglobin content and packed cell volume all decreased but the glucose level increased. The order of decrease in the haematological components of the three fish is in the order of Catla catla > Labeo rohita > Cirrhinus mrigala.     

Influence of marine alga against mercury-induced biochemical changes,structural destabilization,and genotoxicity in marine tiger fish,Therapon jarbua

This study elucidated the protective role of alga against mercury (Hg)-induced toxicity in marine fish. Bath immersion with mercury chloride (HgCl) [0.125 and 0.25?ppm] in combination with the algal extract (3?ppm) to Therapon jarbua offered a significant protection against Hg only induced disturbed liver function, damaged histoarchitecture, elevated oxidative stress, and DNA fragmentation of tissues. Mercury exposure decreased hepatic superoxide dismutase (SOD), catalase (CAT) activities, and the level of nonprotein-soluble thiol (NPSH, GSH), with a concomitant increase in thiobarbituric acid reactive substances (TBARS) in the gill, kidneys, liver, and blood. Algal supplementation diminished the rise in TBARS restoring CAT, SOD, and GSH levels to control. Reduced generation of free radicals may be correlated to protect DNA stability and morphology. This study indicates the ability of alga to ameliorate Hg-mediated injuries. In conclusion, Kappaphycus alvarezii extract did not show any toxicity and its safety is suggestive for using as a supplement in fish food.     

丙烯酰胺对斑马鱼各器官的毒性作用及生殖细胞的DNA损伤

在我国,聚丙烯酰胺(PAM)主要应用于石油开采,其长期暴露于环境中可以降解成有毒的丙烯酰胺(AM)。为探究在降解过程中AM的毒性作用,选择斑马鱼作为受试动物,进行AM长期毒性暴露40 d,考察了对斑马鱼的肝脏、脑组织、心脏、腮等器官的影响情况。结果表明:在2.04 mg·L~(-1)、6.12 mg·L~(-1)和18.36 mg·L~(-1)暴露浓度下,形态学观察斑马鱼的鳃丝,鳃小片和鳃细胞有严重的受损现象;随着浓度的升高斑马鱼肝脏、脑组织和心脏中MDA含量、LDH活力的升高,SDH和Na+-K+-ATPase活力的降低,均对其肝脏、脑组织和心脏造成氧化损伤作用,从而影响了斑马鱼体内细胞能量代谢过程。采用彗星试验检测斑马鱼的生殖腺细胞DNA损伤,结果显示暴露于浓度为2.04 mg·L~(-1)~18.36 mg·L~(-1)的AM后,斑马鱼的DNA损伤均表现为显著差异(P0.01)。上述研究结果均确定了AM对斑马鱼的毒性效应并可造成其生殖腺细胞的DNA损伤。     

Biological weighting of ultraviolet (280–400 nm) induced mortality in marine zooplankton and fish. I. Atlantic cod (Gadus morhua) eggs

In the Gulf of St. Lawrence, Canada, productivity-determining biophysical interactions occur in the upper 0 to 30 m of the water column. The eggs and larvae of several commercially important marine invertebrates and fishes (e.g. Gadus morhua L.) are found in this layer. Measurements of the diffuse attenuation coefficients for ultraviolet-B radiation (280 to 320 nm, UV-B) at various locations in this geographic region indicated maximum 10% depths (the depth to which 10% of the surface energy penetrates at a given wavelength) of 3 to 4 m at a wavelength of 310 nm. This represents a significant percentage of the summer mixed-layer water column: organisms residing in this layer are exposed to UV-B radiation. Laboratory experiments using a Xenon-arc-lamp based solar simulator revealed that cod embryos exposed to UV-B exhibited high wavelength-dependent mortality. The strongest effects occurred under exposures to wavelengths below 312 nm. This susceptibility was also dependent upon developmental stage; mortality was particularly high during gastrulation. At the shorter wavelengths (<305 nm) UV-B-induced mortality was strongly dose-dependent, and not significantly influenced by dose-rate. The biological weighting function (BWF) derived for UV-B-induced mortality in cod eggs is similar to that reported for naked DNA – suggesting that the mortality is a direct result of DNA damage. There was no evidence of a detrimental effect of ultraviolet-A radiation (320 to 400 nm). Calculations based upon the BWF indicate that, under current noon surface irradiance, 50% of cod eggs located at or very near (within 10 cm) the ocean surface will be dead after 42 h of exposure. Under solar spectral irradiance simulating a 20% decrease in ozone layer thickness, this time drops to 32 h. These are first-order estimates based upon surface irradiance taken at a time of day during which the values would be maximal. Nonetheless, they illustrate the relative changes in UV-B impacts that will result from ozone layer depletions expected over the coming decades. It is also important to point out that variability in cloud cover, water quality, and vertical distribution and displacement of cod eggs and larvae within the mixed layer, can all have a greater effect on the flux of UV-B radiation to which fish eggs are exposed than will ozone layer depletion at these latitudes. Received: 2 March 1998 / Accepted: 18 December 1998