Growth and food consumption of 0+ Arctic charr fed pelleted or natural food at six different temperatures

The effects of temperature and diet on the specific growth rate and food consumption of 1-summer-old Arctic charr Salvelinus alpinus were studied. Fish were reared singly in aquaria at six different constant temperatures (5, 9, 13, 16, 18 and 20°C). They were fed Neomysis integer or commercial pelleted food for 2 weeks and growth and food consumption were measured. In both experiments, growth rate increased to an optimum at 15°C. Growth rates were high in the range 13–18°C, with no significant ( P >0·05) differences between temperatures. No significant ( P> 0·05) differences in growth were found between fish at 9 and 20°C. There were no effects of diet on size-adjusted growth rates. The growth efficiency decreased with increasing temperature in both treatments, but the decrease was faster in the Neomysis treatment. Charr seemed to compensate for the high water content (79·5%) of Neomysis by having a higher food intake.     

Aquaculture of fry and fingerling of pike-perch (Stizostedion lucioperca L.) — a short review

This review presents a short survey of pertinent literature on rearing methods of pike-perch fry and fingerlings in ponds, net cages and tanks. The traditional pond culture results in variable numbers of fingerlings, which are sometimes small and are therefore of limited value for stocking. Fingerling production can be increased by manipulating the succession of zooplankton and by stocking adequate quantities of fry in the pond. Culture in illuminated net cages in lakes seems to be a possibility to produce advanced fry. The rearing of larvae in tanks following controlled propagation is initially based on feeding natural plankton. The artificial diet can only be used successfully several weeks later at fingerling size (4–5 cm). High mortality rates of fry and symptoms of food deficiency in liver cells indicate unsatisfactory quality of larval diets tested until now. Lack of an adequate artificial feed considerably hampers further development of intensive rearing of fry. Growth of fry and fingerlings is strongly temperature dependent. Growth is rather poor at 16–18°C and best between 26–30°C. Although there is a short stage of positive phototropism in fry, older pike-perch generally prefer dim light. There is no food uptake at high light intensity and mortalities will occur under such conditions. More research is needed on adequate diets which consider the ontogenetic development of the digestive organs of young fish.     

In situ versus laboratory estimations of length–weight regression and growth rate of Daphnia magna (Branchiopoda, Anomopoda) from an aerated waste stabilization pond

The present paper questions the adequacy of using length–weight regressions and growth rates calculated in the laboratory under constant physico-chemical and food conditions for the estimation of biomass and secondary production of animals living in a variable environment from the physico-chemistry and food availability point of view. Length–weight regressions (LWR) and growth rate of Daphnia magna were determined in situat five key periods of the year. In parallel, LWR and growth rate were determined in laboratory incubators at temperature adjusted to the mean temperature measured during the in situexperiments. LWR estimated from pond daphnids collected during the in situ experiments were, on the whole, not significantly different from LWR established during laboratory experiments, indicating that the food availability was globally similar in the laboratory and in situexperiments, even though food items were substantially different between the experiments. In situ algal biomass was indeed low compared to the algal biomass in laboratory experiments, but high biomasses of bacteria, protozoa and detritus were available for daphnid feeding in the tubes incubated in situ. Growth rate of D. magnawas monitored in situusing 50-ml tubes closed with Nylon net (mesh size = 80 m) and in the laboratory using 50-ml glass flasks. The physico-chemical, bacteriological and algological variables were checked to be similar in the tubes and in the pond. Growth rates varied according to the size of the animal and according to the water temperature. The maximum growth rates were observed for juveniles at 20.2 °C. Growth rates were also determined in the laboratory at temperature corresponding to the mean temperature recorded in the pond during the in situ growth experiments. Differences between in situ and laboratory body length–growth rate regressions (LgR) were observed for the experiments conducted at 15.6 °C and 23.6 °C. Due to differences in LWR and LgR between in situ and laboratory experiments, biomass and daily production estimated from laboratory cultures were found to be significantly, but not severely, higher than biomass and daily production estimated on the basis of in situ experiments. It has been, therefore, concluded that, when the constraints linked to the realization of in situ growth experiments are too strong, the laboratory approach is fully justified.     

Feeding and growth of larval herring,Clupea harengus,in relation to density of copepod nauplii

Synopsis Feeding and growth rates of 1–3 wk old herring larvae from four different stocks were compared in laboratory experiments (8°C). For most of the larval groups, feeding rate was saturated at nauplii (Acartia tonsa, nauplii stages 3–5) densities over 301^–1 (5 g d.w. 1^–1). Specific growth rate increased asymptotically with nauplii density, and reached about 6% d^–1 at densities over 120 nauplii 1^–1. The growth rates attained in the laboratory were similar to field measured growth rates of similarly aged herring larvae at comparable food densities. Since food particles were homogenously distributed in the laboratory tanks, patches of dense plankton concentrations are, thus, apparently not necessary for larval growth and survival in the sea. Growth efficiency differed between larval groups, with large sized larvae being the most efficient in transforming ingested matter into growth. The difference probably relates to different sizes rather than to the different geographical origins of the larvae.     

Density related growth of grass carp, Ctenopharyngodon idella (Val.) in managed small impoundments in Alabama

Growth data for grass carp, Ctenopharyngodon idella , in managed small impoundments were examined for density related effects. Ponds ranged in area from 0.02 to 5.0 ha; various species stocking combinations were used. Growth during the first year of life was strongly affected by density. Total standing stock increased and average size decreased with increasing density. This growth–density relationship was more variable among fish in the second year of life, primarily because of differences in availability of preferred food in different ponds. Survival tended to vary during the first year of life depending on the size at stocking but was uniformly high in the second year.     

The effect of phytoplankton and suspended sediment on the growth of Corbicula fluminea (Bivalvia)

Juvenile Corbicula fluminea Müller (1774) were cultured at 15.3 °C in the laboratory on eight combinations of suspended sediment and phytoplankton. Sediment concentrations were 2.6, 25, 50, and 150 mg l^–1. Chlorophyll a levels were 15.6 and 62.5 µg l^–1. Clam tissue growth was found to be independent of silt concentration but increased at the higher chlorophyll level (p < 0.05). The growth experiment was repeated at 24 °C with chlorophyll a concentrations of 18.9 and 112.6 µg l^–1. Growth was again greater at the higher phytoplankton level (p < 0.05). These results demonstrate that Asiatic clam populations are food-limited most of the growing season in the Northern and Western portions of California's eutrophic Sacramento-San Joaquin Delta where chlorophyll a levels average less than the lower of these values. Comparisons of clam growth in the laboratory and estuary support the food limitation hypothesis as at the higher food concentration laboratory tissue growth was 2.3 and 3.8 times greater during the high and low temperature evaluations than in the estuary.     

Effects of stocking density and food density on survival,growth and yield of laboratory-reared larvae of sea bream Archosargus rhomboidalis (L.) (Sparidae)*

Growth and survival of sea bream, Archosargus rhomboidalis (L.) larvae were affected by both abundance of eggs that were initially stocked in 75 1 rearing systems and by the concentration of copepod nauplii and copepodites that were fed to larvae. Stocking levels were 2, 4, 8,16 or 32 per litre while food abundance was maintained at approximately 100, 500, 1500 or 3000 per litre. Experiments were of 16 days duration at 26° C. Survival was best, often exceeding 60%, when food levels were 1500 or 3000 per litre and when stocking density did not exceed 8 eggs per litre. Growth was best at the lowest stocking densities and highest food levels. The highest total yields in wet weight occurred at 8 per litre stock density and 3000 per litre food level. Mean wet weight per survivor and yield per stocked egg were greatest at the lowest stocking densities and highest food levels. A 500 per litre food level was marginal for growth and survival, and 100 per litre produced significant survival only at the 2 per litre stocking density. Two experiments at 6000 and 10 000 per litre food levels at 4 per litre stock density gave the best observed growth, and survival as good as in any other experiments. Possible relations between sea bream larvae and their food supply in the natural environment are discussed. Results also are discussed in terms of their possible contribution to aquaculture efforts.     

Tradeoffs between respiration and feeding in Sacramento blackfish,Orthodon microlepidotus

Synopsis Suspension-feeding fishes use gill structures for both respiration (lamellae) and food capture (rakers). During hypoxic exposure in eutrophic lakes or poorly circulated sloughs, many fishes, including Sacramento blackfish, Orthodon microlepidotus, increase their gill water flows, in part by increasing ventilatory stroke volumes. Stroke volume increases could compromise particle sieving efficiency by spreading interdigitated gill rakers from adjacent gill arches, although blackfish capture food particles by raker-guided water flows to a sticky buccal root. Using van Dam-type respirometers, blackfish respiratory variables and feeding efficiency (Artemia nauplii) were measured under normoxia (> 130 torr PO₂) and hypoxia (60 torr PO₂). Compared with non-feeding, normoxic conditions, gill ventilation volume, frequency, stroke volume, and gape all increased, while O₂ uptake efficiency decreased, during hypoxia and during feeding. O₂ consumption increased during feeding treatments, and % uptake of nauplii showed no difference between normoxic and hypoxic groups. Thus, blackfish display respiratory adaptations, including increased ventilatory stroke volumes, to survive in hypoxic environments such as Clear Lake, California. Importantly, they have also evolved a particle capture mechanism that allows efficient suspension-feeding under both normoxic and hypoxic conditions.     

Digestibility measurements in juvenile tench [Tinca tinca (L.)] by using a continuous filtration device for fish faeces

Effective pond production of tench [Tinca tinca (L.)] requires the use of formulated diets. However, insufficient information is available regarding the nutrition of this species. Therefore, for the first time, the digestibility of macronutrients by tench was investigated using a sieving method of faeces collection and chromic oxide. A pelleted experimental diet rich in untreated wheat starch revealed the following mean apparent digestibility coefficients (ADCs): dry matter 77.2%, crude protein 89.4%, crude fat 94.3%, ash 30.5%, nitrogen‐free extracts 78.2%, organic matter 83.3% and energy 85.1%. Generally, these ADCs indicate that the ability of tench to digest the macronutrients from pelleted feed correlates with reports of other fish species. However, the tench is more able to digest untreated wheat starch than are carnivorous and coldwater fish species. Compared with carp (Cyprinus carpio L.), the digestibility of untreated wheat starch is less in tench. The faeces sieving method as described in this paper is a useful procedure for further digestibility trials with tench. However, compared with the approach presented here, several adaptations are recommended, e.g. long adaptation times to the rearing facility, high stocking densities, usage of more waterproof food pellets and the addition of formalin to the collection pans.     

The use of whole barley diets fortified with solutions of urea,minerals and vitamins for lambs

For one experiment 45 early-weaned lambs were given one of the following five diets from weaning to slaughter: (1) whole barley with urea, minerals and vitamins added as a concentrated solution; (2) as diet (1) plus 4 g/kg of sodium sulphate in solution; (3) as diet (2) plus 1.2 g of methionine-hydroxyanalogue (MHA)/kg; (4) as diet (2) plus 2.5 ml of 40% formaldehyde added per kg; (5) a control diet containing whole barley and 100 g/kg of a pelleted supplement based on fish meal. Growth rates (g/d) for the five treatments were 218, 253, 253, 256 and 292. Addition of sulphate significantly increased growth rate and food utilization while MHA had no effect; formalin treatment reduced digestibility and food utilization.In a second experiment 58 lambs were used to study the effect of protein supplements for lambs weaned at various ages and weights. Diets similar to (2) and (5) from Expt. 1 were used, while an intermediate diet (6) was made from an equal mixture of diets (2) and (5). As weaning age increased and as live weight at weaning increased, the difference in growth rate and food utilization between lambs receiving diet (2) and those receiving diets (5) and (6) decreased.It is suggested that for most sheep production systems in which concentrates are used either as the sole feed or as supplements, simple fortification of whole grain with the necessary nutrients is all that is required to achieve optimum results.     

Utilization of cyanobacteria by Brachionus calyciflorus: Anabaena flos-aquae (NRC-44-1) as a sole or complementary food source

The rotifer Brachionus calyciflorus can utilize the cyanobacterium Anabaena flos-aquae as either a sole or supplementary food source in laboratory culture. Positive population growth rates accompany food densities of 10 or 100 µg dry weight ml^–1, but slightly negative rates are found at a lower density (1.0 µg ml^–1). These results are consistent for rotifers feeding on two strains of A. flos-aquae, UTEX-1444 and NRC-44-1, with slightly enhanced survivorship and reproduction with the latter food. A 1:1 mixture (by dry weight) of Euglena gracilis and A. flos-aquae (NRC-44-1) produces survivorship comparable to that of control rotifer cohorts fed E. gracilis alone, but elicits significantly greater fecundity and population growth rates than found with the control food suspension at the same biomass density.     

Potential impacts of brown tide, Aureococcus anophagefferens, on juvenile hard clams, Mercenaria mercenaria, in the Coastal Bays of Maryland, USA

The rate of growth of juvenile hard clams, Mercenaria mercenaria, was studied in the Coastal Bays of Maryland during an outbreak of the brown tide, Aureococcus anophagefferens. Brown tide dominated the plankton community during the month of June 2002, with cell densities at several sites reaching category 3 (>200,000 cells ml⁻¹) levels. Temperatures during the bloom were 18.6–27.5 °C. Nutrient conditions preceding and during the bloom were conducive for the proliferation of A. anophagefferens: while inorganic nitrogen and phosphorus were <1 μg at N or P l⁻¹, urea was elevated during bloom development. Organic nitrogen, phosphorus and carbon were in the range of levels observed in previous brown tide blooms and increased following the collapse of the bloom. Growth rates of juvenile clams were significantly lower during the period of the brown tide bloom than following its collapse. Growth rates of M. mercenaria were found to be negatively impacted at brown tide densities as low as 20,000 cells ml⁻¹, or category 1 levels. The low growth rates of M. mercenaria could not be explained by temperature, as the lowest growth rates were found when water temperatures were at levels previously found to be optimal for growth.     

The survival, growth and diet of pike fry, Esox lucius L., stocked at different densities in experimental ponds

Yolk-sac pike fry were stocked at densities of 0.74 – 81.4 m⁻² in two ponds, each divided into eight sectors (mean area 155.8 m²). Growth and survival were recorded from May to December 1985. The growth rates were variable within each sector. The size-range of sampled fish increased throughout the year, but showed no significant correlation with density. Fry survival was initially density-independent but switched by late June/July to density-dependence, ranging from 0.5 to 43.6% of initial stocking numbers. The highest mean daily mortality rates occurred during May-July. The final survival in December ranged between sectors from 0.059 to 11.25% of the starting stock densities. The final biomass per unit area of pike surviving in December was not related to initial stocking density. In Pond 1 the mean biomass produced was 2.21 gm⁻² and in Pond 2 was 3.49gm⁻².
Pike fry < 30 mm fed only on invertebrates; those 30–100 mm took a wide range of invertebrates, cyprinids. sticklebacks and other pike. Cannibalism occurred at most densities between 5.45 and 81.4 fish m⁻².
Where attempts are made to increase pike production in managed populations by releasing small fry, an upper stock density of 5 fry m⁻² is advised if large, density-dependent mortalities are to be avoided.     

Density-dependent influences on feeding and metabolism in a freshwater snail

We used laboratory experiments to assess the degree of, and the underlying mechanism for, density dependence in the grazing rate of the pulmonate gastropod Physella virgata. Both fecal pellet production and uptake and incorporation of ¹⁴C radioisotopes from labeled periphyton were used as indices of grazing rates. Pronounced density-dependent reductions in grazing rate were observed, especially at densities above 4 snails/-25 cm² periphyton grazing area. Radioisotope experiments also indicated that proportions of ingested ¹⁴C periphyton retained in snail tissue and respired as carbon dioxide increased at higher densities, suggesting that both assimilation efficiency and respiratory costs increase at higher densities. Constant replacement of water in aquaria did not remove density-dependent effects on grazing, suggesting that a dissolved metabolite is not responsible. Experiments where tiles were pre-conditioned with snails grazing at several densities actually stimulated grazing in subsequently added snails, suggesting that substrate-borne cues are also not responsible for density-dependent reductions in grazing rate. Behavioral inferference (in the form of shell-shaking after contacts with other snails) did, however, increase at higher densities, and may be partially reponsible for depressed grazing rates.     

Laboratory culture of Chironomus tentans for use in toxicity testing: optimum initial egg-stocking densities

The midge Chironomus tentans Fabricius is a commonly used freshwater invertebrate in sediment toxicity tests. Rigorous laboratory culturing techniques are needed to provide organisms of uniform quality and known age for use in testing and for the continuation of the culture itself. This study was conducted to determine the effect of initial culture stocking density on: (1) post-hatch (larval) dry weight, body length and head-capsule width at 10 and 20 days; (2) time to emergence; (3) number and sex of emergent adults; (4) number of larvae and pupae at test termination (day 42 post hatch); and (5) adult dry weight. Three egg stocking densities were used 690 (1.1 eggs cm^–2), 1043 (1.7 eggs cm^–2) and 1463 (2.4 eggs cm^–2). Mean weight of larvae at 10 days in high density tanks (0.13 mg/organism) was significantly higher (P=0.003) than both the medium and low density tanks (0.10 and 0.09 mg/organism, respectively). No significant differences between the three stocking densities were observed for the body length or head-capsule width at either 10 or 20 days post-hatch. Although not statistically significant, larval dry weight decreased with increased stocking density at day 20. A significantly (P=0.02) greater number of females (173±28) emerged from the low stocking density compared to both the medium and high stocking densities (123±45 and 118±54, respectively). Peak adult emergence for the low and medium stocking densities occurred between days 22 and 25 post-hatch, whereas peak adult emergence occurred between days 30 and 33 for the high stocking density. Survival relative to the initial number of eggs stocked was significantly greater (P=0.007) in the low density treatment compared to that in either the medium or the high density treatments. Mean adult weight exhibited an inverse relationship with initial stocking densities. At test end, there was not a significant difference in the mean number of organisms surviving and emerging in the three density levels. The central tendency for number of organisms surviving for all three treatments was 504 organisms per tank (0.82 organisms cm^–2). The results of this experiment suggest that an optimal egg stocking density of 1.0 egg cm^–2 (600 eggs/tank) be used with the feeding rate identified. This would ensure uniform larvae at the appropriate developmental stage (2nd–3rd instar) needed for toxicological research/testing (e.g. 10 days post-hatch), as well as producing sufficient emergence of males and females for future culture establishment.     

Effects of temperature and food level on growth and development of a planktonic water mite

We analysed the relative effects of food availability and temperature on rates of growth and development of a predatory planktonic water mite, Piona exigua. Growth in length of mites fed Daphnia, Ceriodaphnia and Chydorus was analysed by Gompertz or von Bertalanffy curves; these curves were compared by parallel curve analysis. Growth rates of nymphs and adult female mites increased with temperature; the duration of the imagochrysalis stage decreased. Females grown at 10 °C were smaller at final size than females grown at 15 °C, 18 °C or 22 °C. Females reared at food levels of 15 or 30 prey l⁻¹ grew more slowly and were smaller than those provided with 60 or 120 prey l⁻¹. Nymphs grew more slowly when Daphnia were the only prey, than when smaller prey were available. Food level did not affect nymph growth at 10 °C or 15 °C, but growth at 18 °C or 22 °C may have been slowed at the lowest food levels. Synergistic effects of temperature and food level on nymph growth were apparent only from analysis of growth curves and not from stage duration data.     

Growth,biochemical, fatty acid composition,and mRNA levels of hepatic enzymes in genetically improved farmed tilapia (GIFT,Oreochromis niloticus) (Linnaeus, 1758) at different stocking densities

Investigated were the effects of stocking density on juvenile growth performance, serum biochemistry, fatty acid composition, and mRNA levels of enzymes involved in lipid metabolism in genetically improved farmed tilapia (GIFT), Oreochromis niloticus. Juvenile fish (n = 900) were distributed into 15 tanks at five initial stocking densities (75, 150, 225, 300, and 375 g/m³ per 800‐L tank, represented as D1–D5, respectively). Temperature was maintained at 28 ± 0.3°C, and water flow rate into the tanks was about 8 L/min. The fish accepted floating pelleted feed (crude protein 28%, crude lipid 6%, and gross energy 149.2 KJ/g diet). The feed amount offered was 4%–8% of body weight. At the end of the 60‐day experiment, the final stocking densities were 1,454.18, 2,010.60, 2,913.13, 3,819.73, and 4,380.80 g/m³, respectively. Growth and body mass coefficient of variation were significantly affected (p < .05). No differences in feed conversion rate or survival were found (p > .05). Muscle lipid contents decreased with stocking density from D1 to D5. Serum total protein and cholesterol levels were not affected by the stocking density but the D5 group produced the highest serum cortisol and lactate levels and the lowest glucose levels compared to those of the other groups. Lower muscle saturated fatty acid and n?3 polyunsaturated fatty acid contents were observed in the D4 group than in the other groups, whereas the reverse was found in the n?6 polyunsaturated fatty acid content. Hepatic glucose 6‐phosphate dehydrogenase, fatty acid synthetase, hormone‐sensitive triglyceride lipase, and lipoprotein lipase mRNA levels were significantly upregulated in response to a high density. These data suggest that the GIFT adapted to high‐density conditions by altering the composition of muscle fatty acid and stimulating lipid metabolic enzymes. These mechanisms may help fish respond to high stocking density stress in intensive aquaculture systems.     

Competition for food between an introduced crayfish and two fishes endemic to the Colorado River basin

Synopsis Crayfish are not native to the Colorado River basin (CRB), however they are now established in portions of the mainstem and in many tributaries. I used density manipulation experiments in a laboratory setting to determine intra- and interspecific competition for food between Orconectes virilis, an aggressive polytrophic crayfish now common in the CRB, and two native fishes: Gila chub, Gila intermedia, and flannelmouth sucker, Catostomus latipinnis. I tested each fish species in separate trials. Growth of Gila chub decreased when animal densities increased, however they were more affected by intraspecific competition than by crayfish presence. In contrast, growth of flannelmouth suckers was more affected by crayfish than by intraspecific competition. Crayfish growth was not significantly altered by presence of either fish. Crayfish thus reduced fish growth by competition for food, but the effect differed markedly between the two species. An erratum to this article can be found at .     

Growth and carrageenan quality of <Emphasis Type="Italic">Kappaphycus striatum</Emphasis> var. <Emphasis Type="Italic">sacol</Emphasis> grown at different stocking densities,duration of culture and depth

Kappaphycus striatum var. sacol was grown in two separate studies: (1) at two stocking densities, and (2) at four different depths, each for three different durations of culture (30, 45 and 60 days) in order to determine the growth rate of the seaweed and evaluate the carrageenan content and its molecular weight. The results demonstrated that stocking density, duration of culture and depth significantly (P < 0.01) affected the growth rate, carrageenan content and molecular weight of K. striatum var. sacol. Decreasing growth rate was observed at both stocking densities and at four depths as duration of culture increased. A lower stocking density (500 g m⁻¹line⁻¹) showed a higher growth rate for the shortest durations, i.e. 30 days, as compared to those grown at a higher density. Likewise, decreasing growth rate was observed as depth increased, except at 50 cm after 60 days of culture. A 45-day culture period produced the highest molecular weight at both stocking densities (500 g m⁻¹line⁻¹ = 1,079.5 ± 31.8 kDa, 1,000 g m⁻¹line⁻¹ = 1,167 ± 270.6 kDa). ‘Sacol’ grown for 30 days at 50 cm (1,178 kDa) to 100 cm (1,200 kDa) depth showed the highest values of molecular weight of carrageenan extracted. The results suggested that K. striatum var. sacol is best grown at a stocking density of 500 g m⁻¹line⁻¹, at a depth of 50–100 cm, and for a duration of 30 days in order to provide the highest growth rate, carrageenan content and molecular weight.     

Structure and grazing impact of the protozooplankton community in the waters surrounding the Prince Edward Islands (Southern Ocean)

Fish <1 year old were sampled during 1 year using nets inshore at South Georgia. Some fish were kept in aquaria. Growth rates were estimated using the exponential model. During June to October 1980, field growth rates of Parachaenichthys georgianus and Champsocephalus gunnari were 0.33 and 0.48% SL day⁻¹, respectively. Gobionotothen marionensis (1979 cohort) grew at 0.40% SL day⁻¹ during June to November in the field, and 0.34% SL day⁻¹ in the laboratory from September to March. Notothenia coriiceps grew at 0.28% SL day⁻¹ in the laboratory during September to March. During November to December, Artedidraco mirus grew at 0.82% SL day⁻¹ in the field. The 1980 cohort of G. marionensis grew at 1.39% SL day⁻¹ during November to January in the field. During January, the field growth rate of G. gibberifrons was 1.39% SL day⁻¹. Growth rates increased three-fold from winter to summer. Temperature can only explain one-half of this range in growth rates, whereas all of this range can be explained by food availability. Therefore, seasonal food resource limitation has a major effect on Antarctic fish growth. Received: 30 June 1997 / Accepted: 7 September 1997