Genetic variation and covariation during larval and juvenile growth in Mercenaria mercenaria

Quantitative genetic variances and covariances were estimated for shell length of the hard clam Mercenaria mercenaria (L.) at three larval stages (prodissoconch I, 2 d and 10 d post-fertilization) in 1987 and in 1988 after ca. 9 mo of growth. At each sample interval additive genetic variance was a highly significant component of the total size variation. Narrow sense heritability estimates for shell length ranged between 0.58 (±0.10) for prodissoconch I and 1.08 (±0.29) for 2-d-old larvae. There was significant and positive genetic covariance in prodissoconch I and 2-d larval shell length which resulted in a highly significant genetic correlation (r _g=0.74) between these two traits. This covariance is not surprising since the prodissoconch I comprises the majority of the larval shell of a 2-d-old larvae. The genetic covariances between 2-d-old and 10-d-old larvae and between 10-d-old larvae and 9-mo-old juveniles were low and not significantly different from zero. These results indicate that there is substantial genetic variation for shell growth in M. mercenaria but this variation is not stable during development; the genetic variation in shell growth at one stage of development is not strongly related to the genetic variation in growth during other ontogenetic periods. In this study there were no evident constraints to natural selection for increased shell growth rate during development, which coupled with the high levels of genetic variation may suggest that in nature high rates of larval growth may not be normally subject to significant selective pressure.     

Genetics of larvae and spat growth rate in the oyster Crassostrea virginica

The heritability of oyster (Crassostrea virginica) larval growth rate was estimated to be in the range of 0.25 to 0.50 and a significant part of this genetic variation is of the additive type. Larval growth rate and spat growth rate were found to be highly correlated. These results suggest that a selection program for faster growing larvae and spat would be successful.     

Relation of RNA/DNA ratios to growth for the scallop Euvola (Pecten) ziczac in suspended culture

We examined the relation of RNA/DNA ratios to growth for three size groups of the tropical scallop Euvola ziczac maintained in suspended culture at 8, 21 and 34 m in depth in the Golfo de Cariaco, Venezuel. Various growth parameters indicated that production decreased with depth. This was more likely due to a decrease in seston quality with depth than to temperatures or seston abundance (which were similar at the various depths studied). The RNA/DNA ratio was correlated with the G-index of muscle growth for juveniles (r ²=0.55). A much weaker correlation was observed for the maturing scallops (r ²=0.18), probably because of the interaction between reproductive and somatic growth. In fully mature scallops, somatic growth was negligible and the RNA/DNA ratios appeared to be inversely related to the level of physiological stress of the scallops. Whereas RNA/DNA ratios are difficult to interpret for maturing E. ziczac, because an increased ratio can be due to either increased gonadal or somatic growth, they are useful in predicting growth in juveniles and physiological stress in fully mature scallops.     

Variation in larval life-history traits among reef fishes across the Isthmus of Panama

We tested the hypothesis that regional differences in oceanic productivity have led to the evolution of predictable patterns of regional variation in life-history traits of pelagic larvae of tropical reef fishes. To do so we compared larval traits (egg and hatchling size, larval growth rate and duration, and size at settlement) among closely related reef fishes from the Atlantic and Pacific coasts of the Isthmus of Panama. This comparison provides a control for phylogenetic effects because those regions shared a common fauna prior to the rise of the Isthmus ˜3.5 million years ago, subsequent to which each fauna evolved independently under a very different productivity regime. We measured larval traits of 12 benthic-spawning damselfishes (Pomacentridae: Abudefduf, Chromis and Stegastes) and 13 pelagic-spawning wrasses (Labridae: Bodianus, Halichoeres and Thalassoma). These included members of each genus on each side of the Isthmus and four sets of transisthmian sister species of pomacentrids. Among the pomacentrids we found consistent transisthmian differences in hatchling size, but not in other larval traits. Essentially the reverse pattern occurred among the labrids – larval growth and duration differed consistently among congeners in the two regions, but without consistent differences in hatchling size or size at settlement. Neither relationship is predicted by the regional-productivity hypothesis. Most of the differences were quite small. Stronger phylogenetic effects on larval traits (inter- and intrageneric variation within regions) occur in both families and evidently overwhelm any effect of regional variation in productivity. Reassessment of data that takes into account such phylogenetic effects questions previous conclusions about the existence of regional differences in larval traits among damselfishes in the West Pacific and the Caribbean. Received: 19 January 2000 / Accepted: 26 September 2000     

Ovary development at the onset of gametogenesis is genetically determined and correlated with reproductive traits at maturity in shrimp <Emphasis Type="Italic">Litopenaeus (Penaeus) vannamei</Emphasis>

Besides some mammals and a few model organisms, the presence of genetic variation in ovary or gonad development at ages before that of reproduction has not been established for marine oviparous species nor has the correlation with reproductive traits at maturity. In this investigation, ovary development was evaluated for genetic variability in a full-sib family structured population of subadult shrimp. The numbers of each oocyte type that were present were counted, measured, and used to evaluate the following traits: total oocyte number, mean oocytes diameter, and ovary maturity (OM). The total ovary area was also measured for each female. Genetic variation was assessed through the estimation of the parameter ‘heritability’. The heritability (h ²) of total number of oocytes was zero, but large heritability values were seen for mean oocytes diameter (h ²=0.57±0.27) and OM (h ²=0.71±0.26). Family means correlations between traits in subadults with traits measured in their full-sibs when adults (days to first spawn, total number of spawns, total fecundity, and fecundity at first spawn) indicated the existence of some significant associations between reproductive traits at the two ages. These findings point toward an early genetic determination of reproductive capacity in this crustacean.     

Genetic structure of Dascyllus aruanus populations in French Polynesia

Dascyllus aruanus were collected from 13 different locations in French Polynesia between 15 November 1990 and 15 February 1991 in order to examine larval dispersal on four spatial scales: within-feef, within-island, within-archipelago, among-archipelagoes. Average polymorphism was analysed by protein electrophoresis at two levels (P₉₅=0.285 and P₉₀=0.107) from 12 and 3 polymorphic loci, respectively. Spatial genetic variation displayed a low level of differentiation between populations among archipelagoes, and homogeneity at lower spatial scales. Two hypotheses are proposed to explain the genetic structure observed. The first suggests substantial gene flow between the islands during the pelagic larval phase, the second that the absence of differentiation is the result of recent colonisation. Genetic variation amongst size classes showed a significant heterozygote deficiencies at two loci (PGM ^* and EST-2 ^*) in the smallest size class. This suggests a cyclic selection which affects larvae and adults differentially. The data revealed little differentiation among populations at the different localities, despite the short larval duration of D. aruanus; this suggests that larval duration is not the main factor presently affecting genetic structure in an insular model. Correspondence to: S. Planes at the Université de Perpignan     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin     

Changes in biochemical and fatty acid composition of the razor clam <Emphasis Type="Italic">Solen marginatus</Emphasis> (Solenidae: Bivalvia) during larval development

Aquaculture studies have revealed that polyunsaturated fatty acids are critical for maintaining substantial growth, survival and reproductive rates, and high food conversion efficiencies for a wide variety of marine and freshwater organisms. The aim of this study was to investigate the gross biochemical and fatty acid composition of both neutral and polar lipid compartments of the razor clam Solen marginatus throughout embryonic and larval development. High levels of stored reserves in S. marginatus eggs allow a short larval development, lasting only 8 days. The energy required for embryogenesis was obtained from stored proteins. During larval development from D-shaped veliger until settlement, protein, lipid, and carbohydrate reserves were indistinctly stored for metamorphosis. Although total lipids increased, fatty acids in both neutral and polar lipids decreased during embryonic development. The depots allow a short larval development in which settlement is reached with lower amounts of stored neutral and polar lipids than the contents found in the oocytes. Non-methylene-interrupted dienoic fatty acid levels were similar to those of some polyunsaturated fatty acids, with increasing percentages at the onset of metamorphosis. This study indicates that S. marginatus exhibits a different pattern in the use of gross biochemical and fatty acid reserves during larval development compared to other razor clam and bivalve species, mainly due to the large size of its eggs and the short larval development stage reported in this species.     

Growth and elemental composition (C,N, H) in Inachus dorsettensis (Decapoda: Majidae) larvae reared in the laboratory

Larvae of the spider crab Inachus dorsettensis were reared in the laboratory at constant 12 °C. Development lasted 8 to 10 d in the Zoea I, 10 to 12 d in the Zoea II and 14 to 20 d in the megalopa stage. During this time, larval growth was measured in samples taken every 2 to 4 d as dry wt (W), carbon (C), nitrogen (N), and hydrogen (H); energy content (E) was calculated from C. Biomass and energy (per individual) increased in each larval stage as a parabola-shaped function of age, which could be fitted by a power equation. C, H, and E show a higher percentage gain (relative to the initial values at hatching) than W or N, suggesting that proportionally more lipid than protein is accumulated during larval development. There are cyclical changes in the relative (per unit of W) biomass and energy figures, corresponding to the larval moult cycles: immediately after each ecdysis all these values decrease, presumably due to rapid uptake of water and minerals, then they increase again due to tissue growth and remain high until the next moulting occurs. Cyclical changes in the C/N ratio suggest that proportionally more lipid than protein is accumulated during the initial (postmoult) phase of the moult cycle, followed by a period of balanced or protein-dominated gain during the intermoult and premoult phases. These patterns of growth and elemental composition observed during the complete larval development and in single moult cycles of I. dorsettensis are compared with those described in the literature for other decapod species. This comparison suggests a high degree of similarity in biochemical composition and growth characteristics of larval decapod crustaceans.     

Gpi genotypic effect on quantitative traits in the northern bay scallop,Argopecten irradians irradians

The relationships between glucose-6-phosphate isomerase (Gpi) genotype and quantitative variation related to reproduction and growth were explored over the lifespan of a single cohort of northern bay scallops, Argopecten irradians irradians (Lamarck), from September 1986 to April 1988 in the Niantic River estuary, Connecticut, USA. Analyses revealed that Gpi genotype explained an increasingly significant proportion of variation in scallop size (up to 15%) as the cohort aged. The genotype-specific effects were consistent across sampling dates and among measured traits. There was no evidence for heterosis with respect to size at this locus; however, rare genotypes contributed substantially to the relationship and showed some tendency to fall on the extremes of the phenotypic distribution. The strength and consistency of the genotypic effect on scallop size suggest that genetic variation for Gpi, or some locus in linkage disequilibrium with Gpi, may translate into biochemical and/or physiological variation and affect fitness in this species.     

Maternal allocation of lipid classes and fatty acids with seasonal egg production in Atlantic cod (Gadus morhua) of wild origin

A suite of characteristics is often used to assess egg quality as these properties potentially play important roles in progeny survival and growth. Our objective was to assess egg characteristics including lipid biocomposition of an iteroparous, batch-spawning teleost of wild origin. Maternal allocation to egg number was generally dome-shaped (5 of 8 females) and egg size declined over the breeding season for eight breeding pairs of wild Atlantic cod (Gadus morhua) (n = 43 batches). Egg lipid composition ranged considerably among females and between egg batches within females (e.g., phospholipids 40–86 %; polar 47–87 % and neutral lipids 15–52 % of total lipids; polyunsaturated fatty acids 16–50 % of total fatty acids). Principal component analyses revealed significant inter-relationships among maternal traits, batch sequence and fecundity, and egg size and composition. Seasonal trends with regard to lipid deposition were variable; three females showed consistent declines in lipid parameters (μg egg⁻¹) with both batch number and egg diameter, one female showed consistent increase and the four remaining females showed no trend. The three females that exhibited seasonal declines in egg lipid content were characterized as having high fertilization success (>75 %). Our findings highlight the variability in lipid allocation to eggs of batch spawners of wild origin and characterize the composition of endogenous reserves available during embryogenesis and yolk sac larval stages.     

Effect of temperature on the egg and yolk-sac larval development of common pandora,<Emphasis Type="Italic"> Pagellus erythrinus</Emphasis>

Temperature is one of the most critical environmental factors for fish ontogeny, affecting the developmental rate, survival and phenotypic plasticity in both a species- and stage-specific way. In the present paper we studied the egg and yolk-sac larval development of Pagellus erythrinus under different water temperature conditions, 15°C, 18°C and 21°C for the egg stage and 16°C, 18°C and 21°C for the yolk-sac larval stage. The temperature-independent thermal sum of development was estimated as 555.6 degree-hours above the threshold temperature (the temperature below which development is arrested), i.e. 7°C for the egg and 12.1°C for the yolk-sac larval stage. Higher hatching and survival rates occurred at 18–21°C. At the end of the yolk-sac larval stage, body morphometry differed significantly (p<0.05) between the temperatures tested. The growth rate of the total length increased as temperature rose from 16°C to 18°C, while in the range of 18–21°C it stabilized and was independent of water temperature. The estimated Gompertz growth curve for the yolk-sac larvae of P. erythrinus was (r²=0.992) for the 16°C, (r²=0.991) for the 18°C and (r²=0.981) for the 21°C treatment. The efficiency of vitelline utilization during the yolk-sac larval stage was higher at 18°C.Communicated by O. Kinne, Oldendorf/Luhe     

Growth and content of energy reserves in juvenile sea scallops,Placopecten magellanicus,as a function of swimming frequency and water temperature in the laboratory

The effects of swimming frequency and water temperature on shell growth, tissue mass, and stored energy reserves of juvenile sea scallops, Placopecten magellanicus Gmelin, were examined in a factorial laboratory experiment spanning six weeks in July and August 1992. Individually tagged scallops of similar initial size (22.5±0.1 mm shell height, n=240) were induced to swim to exhaustion at three different swimming frequencies (every day, twice a week, or not at all) in two different water temperature regimes (4 to 7 or 7 to 13°C). The scallops were fed an ad libitum mixture of cultured microalgae. At the end of the experiment, cumulative increase in shell height, dry weight of soft tissues, condition index of dry adductor muscle (adductor muscle dry weight/soft tissue dry weight x 100) and total carbohydrate content of dry adductor muscle were measured for each scallop. Scallops at the higher temperature had significantly greater shell heights, and were in better metabolic condition as evidenced by significantly higher condition indices and muscle carbohydrate contents. The dry soft tissue weights did not differ significantly from their low temperature counterparts. Swimming frequency had no significant effect on shell height, dry tissue weight, or carbohydrate content, but condition index of the adductor muscle increased significantly with swimming frequency. These results show that not only was there no cumulative cost of swimming in terms of shell growth, total soft tissue weight, or carbohydrate content in young scallops, but that condition of adductor muscle tissue was higher in scallops that swam.     

A genetic component to size in queens of the ant,<Emphasis Type="Italic"> Formica truncorum</Emphasis>

The genetic basis of morphological traits in social insects remains largely unexplored. This is even true for individual body size, a key life-history trait. In the social insects, the size of reproductive individuals affects dispersal decisions, so that small size in queens is often associated with reduced dispersal, and large size with long-range dispersal and independent colony founding. Worker size is connected to division of labour when workers specialize in certain tasks according to their size. In many species, variation in worker size has been shown to increase colony performance. In this study, we present the first evidence of an additive genetic component to queen size in ants, using maternal half sib analysis. We also compared intra-colony size variation in colonies with high (queen doubly mated) versus low (queen singly mated) genetic variability. We found a high and significant heritability (h²=0.51) for queen size in one of the two study years, but not in the other. Size variation among queens was greater in colonies headed by a doubly mated queen in one of the study years, but not in the other. This indicates that genetic factors can influence queen size, but that environmental factors may override these under some circumstances. The heritability for worker size was low (h²=0.09) and non-significant. Increased genetic diversity did not increase worker size variation in the colonies. Worker size appeared largely environmentally determined, potentially allowing colonies to adjust worker size ratios to current conditions.Communicated by J. Heinze     

A genetic basis for geographic variation in shell morphology in the bay scallop, Argopecten irradians

The bay scallop, Argopecten irradians, exhibits extensive variation in morphology among geographically separated populations, resulting in the recognition of three major subspecies (A. i. irradians, A. i. concentricus, A. i. amplicostatus). The extent to which morphological variation results from differing environmental conditions is unknown. In the present study, bay scallops from Massachusetts, North Carolina, Florida, and Texas were collected, spawned and the offspring reared in a common garden experiment to determine if scallops cultured under similar environmental conditions exhibited the morphology expected given the geographic origin of their parents. Significant differences among populations were indicated by ANOVA in both the wild-caught (13/14 morphological characters) and cultured (11/14 characters) scallops. Principal components analysis clustered wild-caught scallops according to geographic origin and cultured scallops according to geographic ancestry. The morphological characters most influential in resolving groups were plical width, plical spacing, number of plicae and valve convexity. Geographic variation in morphology apparently has a strong genetic basis, and reflects significant differentiation among disjunct populations of bay scallops. Received: 12 September 1996 / Accepted: 15 October 1996     

Observations on shell deformities,ultrastructure, and increment formation in the bay scallop Argopecten irradians

Shell morphology and ultrastructure were examined in the bay scallop Argopecten irradians, cultured in recirculating seawater systems under various conditions of feeding, lighting, and handling. On a unialgal diet of Thalassiosira pseudonana, scallop growth ranged from 120 to 183 m d^-1 at 20°C in the laboratory, about two-thirds of the growth rate found in the field. However, shell deposited in the laboratory differed from that in the field in several ways. In the field, scallops formed costae as an unpigmented, corrugated marginal shell layer; shell deposited in the laboratory lacked this layer and was therefore darker. Also, microstructure of the exterior shell surface of field scallops was coarsely granular, while that of cultured scallops was relatively smooth. Excessive handling of scallops in the laboratory resulted in marginal thickening of valves, a deformity which was completely arrested by a change from daily to weekly handling. Scallops cultured in the same tank with oysters developed shell-thickening on the interior of the valves. It is postulated that shell abnormalities in bivalves result from disruption of complex behavioral processes associated with shell deposition and may be elicited by a variety of natural and experimental irritants. Under natural lighting regimes and optimal conditions for growth, scallops deposited exactly one shell increment per day, but under continuous lighting, deposition of growth increments often became aphasic. In one 28-d experiment, there was a strong correlation between number of growth increments formed and increase in shell height, suggesting that shell ridge formation occurred intermittently, rather than daily, when shell growth rates fell below approximately 150 m d^-1.     

Accumulation and loss of biomass inLiocarcinus holsatus larvae during growth and exuviation

Liocarcinus holsatus (Fabricius) larvae, of females originating from the Elbe Estuary, FRG, were reared in the laboratory at constant 15°C in May 1988. For each larval stage, developmental time was measured by individual cultures (Zoea I: 6.7±0.7d; Zoea II: 5.0±0.6d; Zoea III: 4.8±0.7 d; Zoea IV: 5.3±0.6d; Zoea V: 6.1±1.1d; Megalopa: 10.45±0.7d). During the entire period of development, dry weight (W), carbon (C), nitrogen (N), and hydrogen (H) were measured daily (Zoea I to V) or every second day (Megalopa). The energy content (E) was estimated from C. Biomass and energy (per individual) increased in each larval stage as a parabolic function of age and is described by power functions. C, H, and E exhibit a higher percentage gain (relative to initial values at the time of hatching) than W and N. It is suggested that proportionally more lipid than protein is accumulated during larval development. Cyclical changes in the relative biomass (% W) correspond to the larval moult cycle, indicating a rapid uptake of water and minerals immediately after hatching and a later increase in tissue growth. Changes in the C:N ratio suggest that during the first period more lipid than protein is accumulated. These patterns of growth and elemental composition are compared with literature data and a high degree of similarity in the growth characteristics of decapod larvae is seen. In addition W, C, N, and H values as well as E were measured for the exuviae of Zoea I to V and Megalopa. The percentage loss of growth rate by exuviae for each larval instar were higher in W (12 to 16%) and C (8 to 12%), and varied between 5 and 10% for N, H, and E.     

Comparison of larval development and growth of the sea cucumberActinopyga echinites: Ovary-induced ova and DTT-induced ova

AdultActinopyga echinites (Jaeger) were collected from northern Taiwan in September 1989. Oocytes were induced to mature by bathing them in ovary juice (ovary-induced ova) or in 10^–2 M dithiothreitol (DTT-induced ova). The percentage of germinal vesicle breakdown (GVBD) increased from 0.4 to 6.4% in the former treatment and to 84% in the latter. After artificial fertilization, the embryos were cultured in seawater (35 S) at 25 to 28°C. Larvae were fed with the algaIsochysis aff.galbana at 10^4–5 cells/ml. Larvae from ovary-induced ova developed faster (18 d to the doliolaria stage) and grew to a larger size (1.13 mm length) than those from DTT-induced ova (20 d to the doliolaria stage and 0.62 mm in length). On the twelfth day, larvae from ovary-induced ova bear lipid spheres. The number of spheres is positively correlated with larval size. Lipid spheres may provide nutrient reserves for larvae during metamorphosis.     

Relationship between larval and juvenile growth rates in two marine gastropods,Crepidula plana and C. fornicata

Previous studies on various marine mollusc species have shown that both larval and juvenile growth rates are substantially heritable, but few workers have examined the extent to which larval and juvenile growth rates covary. We examined the relationship between larval and juvenile growth rates in seven laboratory experiments conducted between 1986 and 1993, using the prosobranch gastropods Crepidula plana Say and C. fornicata (L.). In most experiments larvae were reared individually, measured twice nondestructively to determine larval grwoth rate, allowed or stimulated (daily 5-h exposure to 20 mM excess K⁺ in seawater) to metamophose, and then measured at least twice after metamorphosis to determine juvenile growth rates. Generally, there was no significant (p >0.10) relationship between larval and juvenile growth rates, suggesting that in these two species selection can act independently on the two stages of development. A positive correlation (p=0.007) between larval and juvenile growth rates was observed for C. fornicata in one experiment, but only for offspring from females maturing the most rapidly in laboratory culture. Even for these larvae, however, variation in larval growth rate explained<2% of the variation in juvenile growth rate, so that larval and juvenile growth rates are at most only weakly associated in this species.     

Size-specific changes in amount of lipid and daily growth rate of early larval sardine, Sardinops melanostictus, in the main Kuroshio Current and its offshore waters off eastern Japan

The size-specific nutritional conditions of larval sardines, Sardinops melanostictus, from the main Kuroshio Current and its offshore waters off eastern Japan were assessed by lipid analysis. A rapid lipid analysis technique (diagnostic kits for human serum lipids) was used to measure the different lipid components of individual sardine larvae as indicators of their nutritional condition. Size-specific growth trajectories of individual larvae were estimated by the biological intercept method, and the recent daily growth rate of standard length (SL) was calculated from the 3 d outer increment width on the otolith. Relationships between the amount of larval phospholipid (PL; tissue weight indicator) and SL, and the recent daily growth rate of larva (Gr) and SL, could be expressed by the equations PL = 0.459 SL ^1.77 and Gr = 0.0809 SL − 0.341, respectively. There was no notable difference in these values between the two survey areas. The relationship between the amount of triglyceride (TG) and SL could be expressed allometrically (TG = 0.013 SL ^2.63). The relationship between the index of starvation tolerance (TG/PL) and SL could be expressed by the allometric equation TG/PL = 0.0288 SL ^0.865, suggesting that larger sardine have a higher starvation tolerance than smaller sardine. The TG of the 8 to 9 mm SL size-class larvae in the offshore area was higher than in the main Kuroshio Current. To test whether the TG for each larval sardine in the 8 to 9 mm SL size-class could be correlated with variables (temperature, chlorophyll a and distribution density of the sardine larvae) measured at the sampling stations, correlation analyses were performed. A highly significant negative correlation between TG and distribution density of the sardine larvae was found. A density-dependence effect seemed to influence the fluctuation of the larval storage energy component for short-term needs. Received: 12 March 1998 / Accepted: 26 March 1999