Feeding,conversion efficiencies,and growth of larval mummichogs,Fundulus heteroclitus

Feeding rates, conversion efficiencies and growth of larvae of the mummichog Fundulus heteroclitus, an extremely abundant estuarine fish, were measured at temperatures ranging from 18° to 30°C. The food used was Artemia salina nauplii. At the time of total yolk sac absorption (5 to 7 days after hatching), the feeding rate decreased for a short time, an indication of a shift in metabolism. Higher feeding rates and growth occurred at higher rearing temperatures. The highest conversion efficiency (gross growth efficiency) was 1.1%, at 22°C. Mummichog larvae may be energetically inefficient compared with other fish species, but efficiency might not be critical for this fish, which is an opportunistic omnivore in an energy-rich environment.Contribution No. 291 of the Belle W. Baruch Institute for Marine Biology and Coastal Research, supported by DOE contract No. EY-76-5-09-0869.     

Growth,production and energy transformations in the salt-marsh killifish Fundulus heteroclitus

Populations of Fundulus heteroclitus (L.) at the beginning of a growth season in a New England salt marsh consist of 3 yearly age classes, with the 1-year-old class contributing most of the biomass. Calculation of production rates revealed that the young fish were the most active part of the population and that females were more productive than males. The entire population spent equal amounts of energy in growth and in metabolism. Including the young of the year, we obtained total production of 160 kg dry weight/ha, a value among the highest obtained for natural fish populations. About 5 to 15% of the production is available to predators. Rates of food consumption by F. heteroclitus are high enough to turn over the population of prey relatively often. Popululations of F. heteroclitus are thus capable of exerting an important influence on the abundance and distribution of their prey.Contribution No. 3721 from the Woods Hole Oceanographic Institution.     

Effects of feeding rate on conversion efficiency and chemical composition of the fish Tilapia mossambica

Average daily rations of 14, 35 or 58 mg Tubifex tubifex worm per gram of the fish Tilapia mossambica Peters represent maintenance, optimum or maximum feeding levels. At these levels, conversion efficiency (K ₁) is 5,9 or 24%. An amount of 65 mg worm/g fish/day, when fed under experimental conditions, is converted with the poorest conversion efficiency (4%). Test individuals fed at 11 to 49 mg worm/g fish/day show a decreasing trend in water content (78.1 to 74.8%), and an increasing trend in fat content (32.1 to 44.2%). Below or above this feeding rate range, water content increases, while fat content decreases. The range of individual variations in fat content is nearly 3 times greater than that of ash and 15 times greater than that of water. Test individuals starved for 60 days lose 2.1 mg dry body weight/g/day. This loss is contributed by calorifically equivalent amounts of fat and protein. The endogenous loss of nitrogen by these individuals averaged 0.18 mg N/g body weight/day.     

Tidal and diurnal influence on food consumption of a salt marsh killifish Fundulus heteroclitus

Feeding patterns during four 24-h periods, sampled at 3-h intervals, were investigated for the mummichog Fundulus heteroclitus, in a Delaware, USA tidal marsh. Two factors potentially influencing feeding patterns, time of day and tide height, were examined. On 2 of the sampling periods a low tide occurred in the morning, while on the other 2 sampling periods a high tide occurred in the morning. Results are reported as g-dry wt. of food per g-dry wt. of fish. F. heteroclitus is primarily a daytime feeder that most actively feeds at high tide, regardless of whether or not the high tide inundates marsh surface areas. When tide height was sufficient to inundate the marsh surface, fish invaded these areas and consumed prey characteristic of the marsh surface. F. heteroclitus is an important link in energy transfers between the marsh surface and subtidal systems, enhancing its own energy supplies by consuming marsh surface prey whenever available.     

Intake,digestion, absorption and conversion of food in the fishes Megalops cyprinoides and Ophiocephalus striatus

Different criteria for the assessment of conversion efficiencies of fishes are briefly reviewed, and some important terms defined. In the eurysaline fishes Megalops cyprinoides and Ophiocephalus striatus, transformation of food has been studied as a function of body weight, using both protein and energy transfer criteria. The duration of the 28 experiments conducted varied from 15 to 30 days. Rates of food intake, digestion, absorption and conversion decrease with increasing body weight. Absorption efficiencies are about 97% for the protein fraction, and 91% for the total food consumed; they do not vary appreciably among the individuals of different body weights. Conversion efficiencies of the protein fraction and the total food consumed decrease from about 40% in 2 g individuals to about 15% in 150 g ones. In general, the relations show curvilinear trends, but in weight groups between 10 and 150 g, straight lines are obtained. The slopes of these lines differ in the two species tested. Such differences are related to the physiological state of the fishes and their growth potential. The inverse relation between body weight on the one hand, and food intake, absorption and conversion on the other, are attributed to aging. It is suggested that age affects these intermediate steps via the enzyme-complex of the digestive system. The b values obtained for the relation between body weight on the one hand, and metabolism, food intake and absorption on the other, do not correspond to the surface or weight proportion rule, but to an intermediary type of neither surface nor weight proportionality. As the values obtained are similar, it is concluded that there exists a direct and proportionate relation between metabolism, food intake and absorption. The respective values for the relation between body weight and conversion rate are low indicating that there is a direct, but nonproportionate relation between growth and metabolism.Dedicated to Professor C. P. Gnanamuthu, Madras, on his 66^th birthday, April 29, 1967.     

Nitrogen content of food as an index of absorption efficiency in fishes

Gravimetric estimation of absorption efficiency in fishes is a time-consuming process and still subject to technical errors. Available marker methods require no quantitative recovery of feces, but their applicability is limited (e.g. chromic oxide method) or their reliability is questionable (e.g. Conover's ash ratio method). From over 100 values reported for about 50 fish species, it has been observed that nitrogen content of food holds a positive correlation to absorption efficiency; the relation is significantly (P<0.001) correlated (r=>0.9). Hence absorption efficiency (Ae) of fishes is predictable from the nitrogen content of food with less than 8% error, using the equation log Ae=1.3706+0.5807 log N.Dedicated to Professor O. Kinne on his 61st birthday, August 30, 1984     

Nutritional requirements of the gilthead bream Chrysophrys aurata. Quantitative protein requirements

Six semi-synthetic diets with a casein and corn-starch base, containing 10 to 60% protein were fed for 4 months to young gilthead breams (Chrysophrys aurata, of average initial weight 2.6 g). The object of the study was to determine the optimum amount of protein necessary for growth. The daily intake of the gilthead breams was 5.2 to 2.3% of their live weight. The less protein the diet contained, the more feed the fish consumed. The conversion factor decreased regularly as protein percentage increased. The most favourable conversion rate was 1.4, noted for the group receiving 60% protein. The less protein in the diets, the better the protein efficiency for weight gain and body protein retention. Growth of gilthead breams was proportional to protein content of the diet up to an incorporation rate of 40%. Beyond this, no significant difference in final average individual weight was observed. These results are discussed in relation to those reported for other species of fishes. They show that the gilthead bream requires about 40% protein in its diet, as do most fresh-water species.     

饵料对稀有鮈鲫幼鱼和成鱼生长的影响及摄食优化策略研究

在实验室适宜的饲养条件下,进行不同发育阶段的稀有鮈鲫为期30 d的摄食-生长实验,研究3种饵料在投喂量分别为鱼体湿重的3.0%、4.0%和6.0%的3种摄食水平条件下,对稀有鮈鲫幼鱼(初始湿重为(0.0749±0.02)g)和成鱼(初始湿重为(0.2603±0.06)g)的生长影响。结果表明,不同摄食水平组的稀有鮈鲫幼鱼的相对增重率和特定生长率随摄食水平的增加而增加,但饲料转化率呈下降趋势;而稀有鮈鲫成鱼的相对增重率、特定生长率和饲料转化率的变化趋势基本相同。适宜摄食水平随稀有鮈鲫体重的增加而降低,稀有鮈鲫幼鱼和成鱼的适宜饵料分别为O.S.I.微细颗粒和O.S.I.细颗粒,适宜摄食水平分别为6.0%和4.0%。     

Growth,food utilization and effect of social interaction in the European bass Dicentrarchus labrax

Fingerling and juvenile bass, Dicentrarchus labrax (L.) fed on filleted saithe, Gadus virens, exhibited much higher K ₁ energy and nitrogen utilization efficiencies (growth as % energy or nitrogen intake) than those fed on trout pellets or shore crabs, Carcinus maenas, although the efficiency declined during prolonged feeding. Mean organic absorption efficiency for saithe was 99.5% and was independent of food intake but the efficiency for pellets and crab declined with increasing food intake from 99 to 92%. For each diet the regression of relative growth rate on food intake was linear up to the regression of relative growth rate on food intake was linear up to the highest feeding level and independent of body weight, enabling construction of a simple model for food transformation. The regression coefficient (equivalent to K ₃, the ratio of growth to food consumed in excess of maintenance) for saithe was about 3 times that for pellets in terms of energy and twice that for pellets in terms of nitrogen. The more efficient utilization of saithe for growth and of pellets for maintenance were related to differences in their nutritional value. With each diet, K ₃ and maximum K ₁ for energy were about twice those for nitrogen. Social interaction had a significant effect on food utilization in juvenile bass. Isolated fish showed poor growth and mean K ₁ efficiency in comparison with bass in pairs or groups. K ₃ efficiency and mainternance requirement were similar in individuals and groups, but were significantly greater in pairs. These results are related to behavioural observeations and environmental stress, and the limitations of extrapolating the results of laboratory feeding experiments on fish to other situations is discussed.     

Nitrogen and energy loss via nonfaecal and faecal excretion in the marine teleost Lithognathus lithognathus

Nitrogen excretion and assimilation efficiencies of individual Lithognathus lithognathus (Cuvier 1830), a marine teleost from high energy surf zones in Algoa Bay, South Africa, were determined under laboratory conditions in 1985. Nonfaecal-nitrogen excreted by starved and fed L. lithognathus consists mainly of ammonia with urea and amino acids as secondary excretory products. Ammonia excretion rates were temperature dependant with the excretion rate of starved fish significantly lower than those of fed fish, at all three experimental temperatures. The mass component b of the mass/ammonia excretion equation was temperature independent and ranged from 0.651 to 0.700 and 0.589 to 0.635 for starved and fed fish respectively. The mean percentage of food energy lost via dissolved nonfaecal excretory products (exogenous plus endogenous) was 6.11±6.07%. Assimilation efficiencies ranged from 70.75 to 99.29% for dry matter and from 95.72 to 99.58% on an energy basis. The combined nonfaecal and faecal energy loss was calculated at 11.87% of the ingested energy. The benthic feeding ichthyofauna recycle 255 g total nitrogen per metre strip per year which constitutes 2% of total phytoplankton nitrogen requirements of the surf zone.     

Transformation of food in the fish Megalops cyprinoides

Comparative studies on feeding, digestion, absorption and conversion have been made in the fish Megalops cyprinoides fed with mosquito fish, Gambusia affinis, and prawn, Metapenaeus monoceros, (Pandian, 1967). Feeding as well as digestion rates are faster in individuals fed with Gambusia than in those fed with prawn. The extent of the differences in food intake is more pronounced in smaller individuals than in larger ones. The prawn exoskeleton prolongs the duration of digestion and thus further reduces rate and amount of food intake. Changes in food quality do not affect absorption efficiency. Both, rate and efficiency of food conversion, are higher in M. cyprinoides fed with Gambusia than with prawn.     

Grazing and ingestion rates of nauplii,copepodids and adults of the marine planktonic copepod Calanus helgolandicus

The average grazing and ingestion rates of all stages of the marine planktonic copepod Calanus helgolandicus (Calanoida) from nauplius stage IV to adults were measured experimentally at 15°C in agitated cultures. The chain-forming diatom Lauderia borealis and the unarmoured dinoflagellate Gymnodinium splendens were offered as food. The food concentrations were close to natural conditions and ranged from 36 to 101 g of organic carbon per liter. The medium body weights expressed in g of organic carbon of almost all larval stages raised at 49 g C/1 were identical with the weight of the same stages caught in the Pacific Ocean off La Jolla, California, USA. In a log-log system, grazing and ingestion rates increased almost linearly with increasing body weight. Grazing rates ranged from 4 to 21 ml/day/nauplius stage IV to 286 ml to 773 ml/day/female. Ingestion rates increased from 0.2 g to 0.8 g C/day/nauplius stage IV to 18 g to 69 g C/day/female. Grazing and ingestion rates per unit body weight decreased gradually with increasing body weight. The daily ingested amount of food decreased from 292 to 481% of the body weight (g C) of nauplius stage V to 28–85% of the body weight of adult females. Grazing and ingestion performances of all stages increased with increasing particle size. Grazing rates decreased and ingestion rates increased with increasing food concentrations. The published data on food intake of the different age groups of C. helgolandicus show that the young stages of herbivorous planktonic copepods can play a major part in the consumption of phytoplankton in the sea due to their high grazing and ingestion rates.     

A comparison of absorption and assimilation efficiencies between four species of shallow- and deep-living fishes

We captured two species of deep-sea zoarcids, Melanostigma pammelas and Lycodapus mandibularis, from Monterey Bay California and maintained them in the laboratory. One shallow-water zoarcid, Eucryphycus californicus, and an ecologically and morphologically similar stichaeid fish Xiphister atropurpureus were collected from intertidal and subtidal habitats in Monterey Bay. We investigated the absorption and assimilation efficiencies of these fishes to determine whether deep-sea species have evolved mechanisms to increase their efficiency of food use. Fishes were placed in experimental chambers and fed a known quantity of food. Ammonia excretion was measured and feces were collected daily. Both food and feces were analyzed for water, protein, lipid and ash to determine specific absorption efficiencies. Absorption ranged from 87.7 to 92.4% and assimilation efficiencies from 84.0 to 86.5%. Meal sizes from 0.5 to 4.0% of body weight did not affect the results. No significant differences were found between deep-sea and shallow-water species fed single meals or fed ad libitum for 10 days. This suggests that the selective pressure to maximize absorption and assimilation is universal and is not affected by the productivity of the habitat occupied. However, the relative size of the intestine in the deep-sea species was significantly smaller suggesting that they had a lower metabolic cost to maintain their digestive apparatus. It could not be concluded whether this was the result of pressure to conserve energy or rather the tendency of the shallow-living species to ingest more refractory material (i.e. sediment, algae).     

Trophic ecology and fecundity of Fundulus heteroclitus in Chezzetcook Inlet,Nova Scotia

The ontogenetic and seasonal dietary shifts of the mummichog Fundulus heteroclitus (L.), from eelgrass (Zostera marina L.) beds off the shore of Halifax County, Nova Scotia, Canada were examined over a 5 mo period in 1980. Switches to abundant, stationary, or high-calorie prey types were observed in the foraging period. Smaller fish generally took less diverse prey. The indices of foregut fullness varied according to body length and season. The successive 10 mm total-length classes of the mummichog (14 to 101 mm TL) consumed whole prey of increasing size. In comparison with populations living further south, the Nova Scotia fish had a shorter spawning period, lower fecundity, and smaller eggs.     

Assimilation efficiency of a temperate-zone intertidal fish (Cebidichthys violaceus) fed diets of macroalgae

We studied assimilation efficiencies of the temperate-zone intertidal fish Cebidichthys violaceus (Girard, 1854) fed in the laboratory on each of the following species of macroalgae: Spongomorpha coalita (Chlorophyta), Ulva lobata (Chlorophyta), Iridaea flaccida (Rhodophyta) and Porphyra perforata (Rhodophyta). Together, these 4 algae make up over 75% of the natural summer diet of C. violaceus. Assimilation efficiency was calculated by proximate organic analysis of food and feces; the amount of ash in food and feces was used as a standard. Depending on the algal species, the fish assimilated 43 to 81% of the protein, 21 to 44% of the lipid, 45 to 62% of the carbohydrate and 31 to 52% of all three classes of organic material combined. These data are the first results showing that a temperate-zone marine fish can assimilate macroalgal constituents. Protein, carbohydrate and total organic material were absorbed more efficiently from rhodophytes than from chlorophytes. Conversely, lipid was absorbed more efficiently from chlorophytes than from rhodophytes. These results are compared with previous work showing that C. violaceus in nature eats more chlorophytes than rhodophytes, but in laboratory preference tests prefers rhodophytes to chlorophytes.     

Sources of cyclopropanoid fatty acids in the mummichog Fundulus heteroclitus

The saturated fatty acids in lipids of the mummichog Fundulus heteroclitus (L.) contain the characteristic bacterial cis-9,10-methylenehexadecanoic and cis-9,10-methyleneoctadecanoic acids in addition to straight-chain, iso, anteiso and multi-branched fatty acids. To ascertain whether the cyclopropanoid fatty acids (CFA) might be used as a marker for bacterial components in the mummichog diet, the saturated fatty acids of detritus were compared with those of fish intestines and their contents, and with those of eviscerated fish. The relative amounts of individual fatty acids in each sample indicated that microorganisms or invertebrate animals that consume microorganisms constitute a part of the mummichog's diet. A similar analysis of anterior and posterior sections of the fish intestines showed a higher concentration of CFA in the posterior section, suggesting that at least some of the cyclic acids might originate from the resident bacterial gut population. When a group of mummichogs was maintained on a beef-liver diet devoid of cyclic acids, carcass lipids were not depleted of CFA. Instead, an increase in concentration was observed. It was concluded that the intestinal microflora have a role in supplying CFA to mummichog lipids. Since the overall results indicate that these acids are derived from commensal bacteria as well as from bacteria associated with the diet, it is not possible to use such compounds as food-chain markers.     

Altered c-Fos expression demonstrates neuronal stress in mummichog, <Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>, exposed to <Emphasis Type="Italic">Pfiesteria shumwayae</Emphasis> and <Emphasis Type="Italic">Chaetoceros concavicornis</Emphasis>

To better understand sublethal effects of harmful algal blooms (HABs) on fish, mummichog, Fundulus heteroclitus (L.), were exposed in the laboratory to varying, environmentally relevant densities of Pfiesteria shumwayae (Glasgow et Burkholder, CCMP 2089, dinoflagellate) and Chaetoceros concavicornis (Mangin, CCMP 169, diatom). Two experiments were conducted during the spring of 2003 and 2004 to quantitatively examine the effects of acute (2 h) P. shumwayae and C. concavicornis algal exposure on mummichog brain activity using c-Fos expression as a marker of altered neuronal activity. Brains from HAB-exposed fish were removed, sectioned, and stained using immunocytochemistry prior to quantifying neuronal c-Fos expression. Fish exposed to P. shumwayae and C. concavicornis showed increased c-Fos expression compared to unexposed control fish. A significant dose-response relationship was observed, with increased labeling in brains of fish exposed to higher cell densities for both HAB species tested (P ≤ 0.01). Increased labeling was found in the telencephalon, optic lobes, midbrain, and portions of the medulla. The greatest increases in expression were observed in the telencephalon of P. shumwayae-exposed fish, and in the telencephalon and optic lobes of C. concavicornis-exposed fish (P ≤ 0.01). These increases in c-Fos expression are consistent with other physical and chemical stress exposures observed in fish. Neuronal stress, evidenced by c-Fos expression, demonstrates a sublethal effect of exposure and changes in brain activity in fish exposed to HAB species.     

Filtering rates of the juvenile Atlantic menhaden Brevoortia tyrannus (Pisces: Clupeidae), with consideration of the effects of detritus and swimming speed

From July to September 1982 feeding experiments were conducted with 138-mm fork length Atlantic menhaden Brevoortia tyrannus (Latrobe) (Pisces: Clupeidae) to determine their particle size-specific feeding abilities. Monoculture clearing-rate experiments showed that the minimum size of particles filtered, the minimum size threshold, for 138-mm fish is 7 to 9 m. Filtration efficiency for three species of phytoplankton below the minimum size threshold. Pseudoisochrysis paradoxa, Monochrysis lutheri, and Isochrysis galbana, averaged 1.0% (n=14). Tetraselmis suecica, Prorocentrum minimum, and 2-celled Skeleionema costatum, phytoplankton which are larger than the minimum size threshold and smaller than the 20-m upper limit for nanoplankton, were filtered at efficiencies averaging 21% (n=24). S. costatum chains of 3 to 6 cells, prey particles exceeding the size limits of nanoplankton, were filtered at average efficiencies ranging from 22 to 84%. The mean filtration efficiency for Artemia sp. nauplii (San Francisco Bay Brand) of 36% (n=7) was lower than for smaller phytoplankton prey. The presence of detritus at concentrations usually encountered in nature enhanced filtering efficiency and lowered minimum size thresholds at which phytoplankton were retained. For small food particles, filtering efficiency decreased as swimming speed of the menhaden increased. As menhaden grow, their feeding tepertoire shifts to larger planktonic organisms.Contribution No. 1201 Virginia Institute of Marine Science     

An ecological simulation reveals balancing selection acting on development rate in the teleost Fundulus heteroclitus

Semi-lunar spawning in the intertidal zone has been observed in many teleost species. Because the eggs of these species are placed at elevations reached only by the highest tides, the timing of hatching is critical to survival. To quantify and clarify the relationship between development rate, hatching, and survival we developed a computer model that simulated the reproductive strategy in one such species, Fundulus heteroclitus (Linnaeus). We determined expected hatching success for F. heteroclitus embryos as a function of development rate by simulating spawning, embryo development, and hatching of F. heteroclitus embryos in Delaware Bay. Our simulation analysis included manipulation of several model parameters. The results indicated that there is balancing selection acting on development rate in this species. In particular, false hatching cues (rainfall) increased mortality among embryos with very fast and very slow development rates. The simulation results did not appear susceptible to manipulation of the sensitivity of embryos to false hatching cues. The timing of spawning relative to the spring tides determines which development rate is optimal in terms of average hatching success. Including a constant daily mortality rate in our simulations, as a simple model of predation and/or disease, shifted the optimized development rates to faster values. These results may be applicable to other fish species with similar reproductive strategies. Received: 2 September 1996 / Accepted: 17 September 1996     

The importance of an infrequently flooded intertidal marsh surface as an energy source for the mummichog Fundulus heteroclitus: An experimental approach

Fundulus heteroclitus is known to ascend onto the marsh surface to feed. Our study investigated whether the marsh surface food items are a necessary source of caloric intake for the Canary Creek, Delaware, USA population of this species. Enclosure techniques were used to restrict mummichogs from the marsh surface and the growth rates of these fish were compared to those having access to the marsh surface. Growth rates were significantly higher for mummichogs allowed access to the marsh surface. Food addition and density reduction experiments showed that food availability per fish, rather than behavioral responses due to fish crowding, was responsible for the increased growth. Although food was available in the subtidal portion of the habitat, it was of insufficient quantity for fish at natural density to grow at a normal rate, and mummichogs must utilize the marsh surface for at least a portion of their energy intake.