Carbohydrate metabolism of two crustaceans during starvation

In Emerita asiatica and Ligia exotica, starvation for 1 week steadily depleted the free sugar values, indicating their involvement in nutritive physiology. Protein-bound blood sugars showed no change during starvation, indicating their non-involvement in general carbohydrate metabolism; it is suggested that they may be involved in the moulting physiology related to the synthesis of structural components such as chitin. The extent of glycogen storage is found to be greater in the anomuran E. asiatica than in the isopod L. exotica. In the latter, the fall in free sugar values during starvation is greater, due to meagre storage of glycogen in the hepatopancreas. these features are related to habitat and availability of food in the environment.     

Blood sugar metabolism during moulting in the isopod crustacean Ligia exotica

Blood sugar metabolism of the isopod Ligia exotica Roux was studied in relation to chitin synthesis during the moult cycle. The blood sugars were found to occur free, as well as bound with blood proteins, during all moult stages, as in Emerita asiatica. The variety of free blood sugars is smaller in L. exotica; it comprises only glucose and glucose-6-phosphate. The protein-bound blood sugars are glucose and glucosamine. The blood-volume values do not differ markedly during the moult cycle. Estimations of blood sugars showed a marked rise of sugar levels in the blood during the premoult phase, and a subsequent fall during the freshmoult phase, when chitin is actively synthesised. The chitin values also showed a definite correlation with the blood-sugar values. The increased resorption of chitin during the premoult stage, and its liberation into the blood stream as glucosamine, is indicative of its reutilisation in chitin synthesis as in E. asiatica and the genus Cecropia. As more immediate precursors of chitin, such as uridine diphosphate and uridine diphophate acetylglucosamine compounds, were absent in the blood, a thorough biochemical study of the epidermis is suggested to locate the intermediate precursors of chitin which are possibly derived from blood sugars.     

Blood sugars in relation to chitin synthesis during cuticle formation in Emerita asiatica

Blood sugars of the crustacean Emerita asiatica were studied during the moult cycle to examine their possible role in the synthesis of chitin, which takes place at the time of moulting. Blood sugars occur both in free state as well as bound with proteins, and show quantitative fluctuations in different phases of the moult cycle. The values of sugars were estimated in relation to total blood volume, to obviate the effects o absorption of water and consequent dilution of blood taking place during moulting. The data obtained suggest that the presence of glucosamine only during premoult stage may be due to resorption of chitin preparatory to the shedding of the cuticle. The fluctuations in the protein-bound blood sugars appear significant in the context of the synthesis of chitin, in view of the correlation seen between these and the chitin content. The above suggestion is supported by the observation that they are not involved in the nutritive metabolism of the crustacean. The protein-bound sugars do not appear to undergo further changes in the blood, as may be inferred by the absence of uridine diphosphate, and uridine diphosphate acetylglucosamine compounds, which have been suggested to be the more immediate precursors of chitin during the moult cycle of insects. It is suggested that protein-bound sugars may be transported, as such, to the epidermis, which may be the site of the final steps in chitin synthesis.     

Change in the gross biochemistry and mineral content accompanying the moult cycle in the Antarctic krill Euphausia superba

Concentrations of water, ash, protein, chitin, lipid, calcium, magnesium, sodium, potassium, strontium and copper were measured in individuals from a laboratory population of Antarctic krill, Euphausia superba Dana, over the course of a moult cycle. Significant changes in all variables were encountered. Total ash, lipid, calcium, magnesium and strontium all increased in concentration following moulting. Water, protein and copper concentrations all decreased following ecdysis and increased again towards the end of the moult cycle. The major ions sodium and potassium fluctuated around mean levels. Cast moults of E. superba were shown to be a drain on the ionic load of the krill, and the losses inherent in exuviation could account for much of the variation observed during the moult cycle.     

Endocrine regulation of carbohydrate metabolism during the moult cycle in crustaceans I. Effect of eyestalk removal in Ocypoda platytarsis

The endocrine regulation of carbohydrate metabolism in the decapod Ocypoda platytarsis was investigated by eyestalk ablation at all stages of the moult cycle. Eyestalk removal (and hence removal of a substance probably produced by it) profoundly influences the carbohydrate level of the major storage organ, the hepatopancreas. The changes taking place storage organ, the hepatopancreas. The changes taking place in hepatopancreatic glycogen are reflected in the blood sugar (glycemic) level. The blood-sugar level is also related to dynamic events taking place in the cuticle during the moult cycle. At the intermoult and postmoult stages, the eyestalk factor directs the sugar supply for energy metabolism, and prevents sugar deposition as glycogen in the hepatopancreas. At the premoult stage, the activity of the eyestalk is suppressed by the moulting gland (the Y organ), as eyestalk ablation does not alter the blood-sugar value or the hepatopancreatic glycogen-level. After the increase at the freshmoult stage due to eyestalk ablation, the blood-sugar level tends to re-attain its previous low level at the postmoult stage.     

Two distinct forms of the chitin-degrading enzyme N-acetyl-β-d-glucosaminidase in the Antarctic krill: specialists in digestion and moult

In the Antarctic krill Euphausia superba two forms of the chitinolytic enzyme N-acetyl-β-d-glucosaminidase (NAGase, EC 3.2.1.52) have been described, previously identified as NAGase B and NAGase C. Here, we demonstrate the organ-specific distribution and physiological relevance of both forms using a polyclonal antibody preparation which allows them to be distinguished immunologically. While NAGase B was localized in the integument and displayed a pattern of activity related to the moult cycle, the activity of NAGase C was independent of the moult cycle and was predominantly found in the gastrointestinal tract. Accordingly, NAGase B played a significant role in chitin degradation during the krill's moult, whereas NAGase C participated in the digestion of chitin-containing dietary components. Chromatographic elution profiles of isolated organs confirmed the immunological results by displaying characteristic organ-specific patterns in NAGase activity. The molecular characteristics of the moulting form, NAGase B, may further indicate a vesicular transport of moulting enzymes from the epidermis into the ecdysial space. Based on our results we develop a hypothesis explaining the concurrent processes of simultaneous chitin degradation and chitin synthesis occurring during moult. Received: 30 December 1998 / Accepted: 23 April 1999     

Observations on the biochemical changes in gonads and other organs of Uca annulipes,Portunus pelagicus and Metapenaeus affinis (Decapoda: Crustacea) during the reproductive cycle

The fluctuations in biochemical constituents such as water, nitrogen, non-protein nitrogen, protein, lipid and glycogen in gonad, muscle and hepatopancreas have been followed in 3 decapod crustaceans, Uca annulipes (Latreille), Portunus pelagicus (Linnaeus) and Metapenaeus affinis (Milne-Edwards). The water and ash content of the entire body show no systematic fluctuation in relation to the annual reproductive cycle. The water content of the ovary diminishes as it matures. In the ovary of these crustaceans, the lipid fluctuated greatly in relation to the reproductive cycle. The maturing ovary contains more lipid than an immature or spent ovary per unit tissue weight. The changes in the biochemical constituents in the testis are not so pronounced as in the ovary, since the testicular cycle is often drawn-out and almost continuous in these crustaceans. There is an inverse relationship between water content and lipid content of the hepatopancreas; the greater the fat content, the lesser the water content. The hepatopancreas in these crustaceans is apparently a storage organ and contains much lipid and glycogen. At the height of the breeding season, when gonad production is intense, there is an indication of the mobilisation of at least a part of the lipid from hepatopancreas to gonad.Dissertation for the award of a Doctorate Degree of the Kerala University.Formerly the Marine Biological Laboratory.     

A refinement of the moult-staging technique for Antarctic krill (Euphasia superba)

The moult-staging technique for euphausiids developed by Buchholz was examined experimentally to obtain estimates of the relative duration of each of the constituent stages. The method entailed removing uropods from indiviually maintainedEuphausia superba Dana on the day of moulting and for each day post-moult, and examining them microscopically to determine the moult stage. The krill were kept alive following uropod removal until the next moult so that the stage observed in the excised uropod could be referenced to both ends of the moult cycle for each individual examined. The proportion of the moult cycle occupied by each stage at 0°C was thereby calculated.     

Ovarian development and spawning in relation to the moult cycle in Northern krill, Meganyctiphanes norvegica (Crustacea: Euphausiacea), along a climatic gradient

Adaptive processes linked to reproduction were studied comparatively for three populations of Northern krill, Meganyctiphanes norvegica (M. Sars, 1857), sampled during winter and summer cruises in the Clyde Sea (W Scotland), the Kattegat (E Denmark), and the Ligurian Sea (NW Mediterranean). The aim was to investigate the functional relationship between egg production and moulting under contrasted climatic and environmental conditions. A staging system for female sexual development established for live krill was complemented by a histological study of the ovary at various developmental steps. During the reproductive season, all adult female krill were engaged in cyclical egg production. During experiments, female krill released one batch of mature oocytes in one or two spawning events. The ovary of postspawn female krill still contained developing oocytes for another egg batch. In the non-reproductive period, all female krill had a resting ovary. Ovarian structure and pattern of egg production were identical in the three populations, but seasonal timing of egg production was different. The model proposed for the Ligurian population of the annual cycle of ovarian development can be extended to the other two populations, taking into account the seasonal characteristics of each site. Random field samples were staged simultaneously for moult cycle and for sexual development. Moult stages and the seasonal variation of the intermoult period were studied for the Kattegat population using multi-year data and compared to data obtained during summer/winter cruises in the Clyde and the Ligurian Sea. At the three sites, intermoult period was shorter and temperature-dependent during the reproductive period, concurrent with the season of greatest food availability. During most of the year and the period of sexual rest, moulting activity was reduced. The relationship between spawning and the moult cycle was studied comparatively for the three populations. Eggs were released during the premoult phase of a “spawning moult cycle”, in one or two spawnings associated with apolysis and Moult Stage D1, respectively. Yolk accumulation for the next egg batch was completed during an alternating “vitellogenic moult cycle”. A model for the timing of cyclical egg production in relation to moulting, as proposed for the Kattegat, can be extended to the other populations, taking into account intermoult period variation with temperature. Temperature appeared to be the principal environmental factor controlling growth (through moulting) and egg production during the reproductive season, in connection with favourable trophic conditions. Received: 22 December 1997 / Accepted: 29 August 1998     

Energetics,moult cycle and ecdysteroid titers in spider crab (Hyas araneus) larvae starved after the D0 threshold

Zoea I larvae of Hyas araneus L. (Decapoda: Brachyura: Majidae) were dredged in January 1986 from the German Bight and reared in the laboratory at constant 12°C, until they reached the transition of stages C/D₀ of the moult cycle (4 d after hatching). This developmental stage had previously been found to correspond with the point of reserve saturation (PRS) which allows autonomous (food-independent) development through the rest of the moult cycle and hence, was termed the D₀ threshold. One part of the larvae was continually fed (control), another group was starved from the D₀ threshold until moulting to the zoea II instar. In these two experimental groups, as well as in the two groups of zoea II larvae obtained from the different feeding conditions, the course of the moult cycle, biomass (dry weight, W; carbon, C; nitrogen, N; hydrogen, H; energy, E; the latter estimated from C), and ecdysteroid titers (measured with a radio-immuno-assay as ecdysone equivalents) were investigated. When the larvae reached the PRS, they had gained 90% in W, 72% in C, 32% in N, 53% in H, and 65% in E, since hatching, corresponding to an accumulation of 87% of final W and 62 to 69% of C, N, and H reached later, at the end of the mould cycle in the control. The period of starvation caused a 2.5-d delay of the moult cycle, mainly in late premoult, and significant losses of biomass and energy. Starved and fed larvae secreted similar amounts of moulting hormone per individual, but with a reduced rate in the starved group, thus causing developmental delay. Zoea II larvae moulting after starvation contained less than half of the control biomass and energy, and even less than a freshly hatched zoea I. Growth rate was only slightly enhanced in these zoea II larvae as compared to the fed control, but losses of biomass, mainly of lipids, were partly compensated by a 4-d prolongation of their moult cycle, chiefly (3 d) in stage C. Biomass curves were almost parallel in the two experimental groups of zoea II larvae, with significantly higher values in the control during all stages of the moult cycle. However, similar relative proportions (74 to 89%) of late premoult biomass and energy were reached at the D₀ threshold, regardless of different feeding history and initial or final values in a given group. The ecdysteroid titer curve of the zoea II which had moulted from starved zoea I was very similar to that in control larvae, but with a 3-d delay in the occurrence of premoult peak concentration (in both groups in stage D₁). Regulation and coordination of moult cycle, ecdysteroid titers, and growth in the larval development of decapod crustaceans are discussed, with special reference to the D₀ threshold.     

Functional cytology of the hepatopancreas of Penaeus semisulcatus (Crustacea: Decapoda) during the moult cycle

In the shrimp Penaeus semisulcatus, the digestive gland functions as the main site for the digestion, absorption and storage of nutrients and metals during the feeding stages of intermoult and for the mobilization of reserves during the non-feeding stages of pre-and post-moult. These different activities are reflected directly in the cytology of the gland. All the epithelial cells show marked fluctuations in frequency and changes in structure, form and function which occur in phase with the moult cycle. The shrimp were collected offshore from Kuwait in 1980.     

R-cells and the digestive cycle in Penaeus semisulcatus (Crustacea: Decapoda)

Shrimp, Penaeus semisulcatus, were collected from Kuwaiti waters in 1980, and the R-cells in the hepatopancreas were observed during the digestive cycle. During the first 12 h they absorb soluble material from the lumen of the gland and, subsequently, they take-up soluble and particulate material from the haemolymph. Within the cytoplasm, apical and basal systems of smooth endoplasmic reticulum are concerned with these processes. Pinocytosis only occurs at the basal cell membrane. The absorbed material is stored as lipid and glycogen, and copper, sulphur and other elements are accumulated in a large dense vacuole. These activities are integrated with those of the other cell types to effect the assimilation of food in the hepatopancreas. A scheme is proposed which accounts for the main processes of digestion. The hepatopancreas governs the duration of the digestive cycle, which takes at least 24 h. Observations on the nature of intracellular storage and waste products could be used to assess the dietary requirements of artificially reared shrimp.     

Sediment texture and cannibalism affect survival during moult in Saduria entomon (Isopoda)

Laboratory experiments were performed to investigate the effects of sediment texture and presence of non-moulting conspecifics on size-specific moulting survival of the omnivorous benthic isopod Saduria entomon (L.). The moult survival of S. entomon was significantly higher in coarse sand than in very soft muddy clay sediment. However, there was no size-related difference in moult survival between the substrates. Due to cannibalism, moult survival of S. entomon was substantially reduced in the presence of non-moulting conspecifics. Moult survival was higher when the moulter was larger than the non-moulters compared to when the moulter was smaller. In contrast, the absolute size of the moulter or the non-moulters did not affect moult survival. The sex of the moulting or non-moulting isopods had no effect on the survival of the moulters. Based on the results from laboratory experiments, sediment texture and cannibalism are suggested to affect survival during moult of S. entomon in the field. Received: 12 February 1998 / Accepted: 14 November 1998     

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.     

Growth of the scyllarid lobsters Ibacus peronii and I. chacei

The growth rates of the morphologically similar scyllarid lobsters Ibacus peronii (Leach, 1815) and I. chacei (Brown and Holthuis, 1998) are described using data from a tag/recapture study and from tagged lobsters kept in captivity. Within particular size classes, we found no differences in moult increments between male and female I. peronii nor between male and female I. chacei. Small individuals of both species always had larger moult increments than larger individuals. For I. peronii, females moulted more frequently than males, and smaller size classes moulted more frequently than larger size classes. Female I. peronii therefore grew more quickly than males and reached their estimated size at sexual maturity (51 mm carapace length) after ∼2 yr. Moulting of I. peronii was seasonal, with most lobsters (96.3%) moulting between October and January. We found no differences in growth rates of I. peronii at two locations along the east coast of Australia: Coffs Harbour in New South Wales (30°18′S; 153°08′E), and Lakes Entrance in Victoria (37°53′S; 148°00′E). For I. chacei, we found no differences in the frequency of moulting between males and females and, because we also found no differences in the moult increments between males and females, the growth rates of both sexes were the same. Received: 14 August 1999 / Accepted: 20 January 2000     

Lipid-class composition of organs and tissues of the tiger prawnPanaeus esculentus during the moulting cycle and during starvation

Neutral lipid and phospholipid fractions and their component lipid classes in the digestive gland, abdominal muscle, epidermis and cuticle ofPenaeus esculentus Haswell were analysed and compared during the moulting cycle and during starvation. The prawns were collected from Moreton Bay, Queensland, Australia, by trawling during 1985–1987, and were fed on a standard, semi-purified diet. The digestive gland appears to be a major site of lipid synthesis, storage and mobilisation in preparation for moulting. Neutral lipid, 59 to 80% of which was triacylglycerol, was the larger fraction. It accumulated during early premoult, mainly due to the increase in triacylglycerol. The digestive gland contained only 18% of the total body lipid, or 8% of body lipid as triacylglycerol. Thus, the reserve lipid available for energy production is very small. Digestive gland triacylglycerol was markedly depleted after 4 d starvation and was almost completely absent after 8 d. In the other tissues, the major fraction was phospholipid, of which over 50% was phosphatidylcholine and up to 20% phoshatidylethanolamine; cholesterol was the major class in the neutral lipid fraction and appeared to be very stable. Most of this lipid was probably a component of cellular membranes. The lipid composition of muscle changed very little during the moulting cycle: total lipid levels in the epidermis were high in late premoult and early postmoult, when new cuticle is being secreted, but the proportions of the component lipids were closely similar. Cuticle lipid, together with other major components, was resorbed from the old cuticle prior to ecdysis, but the cuticle phospholipids appeared to be labile at all moult stages. The total of all lipids in fedP. esculentus was about 3.6% dry weight, of which about 70% was phospholipid. Earlier research had shown that when digestive gland lipid is exhausted after a short period of starvation, muscle is metabolised for energy. The present research showed that in the remaining muscle only about 13% of lipid was lost after 21 d starvation, mostly as phosphatidylcholine. This is in keeping with the need to maintain this tissue in a functional state. In contrast, epidermal lipid levels were markedly reduced after only 4 d starvation and the proportions of phospholipids changed significantly. This sensitivity of the cuticle lipids to starvation may be the cause of delayed moulting, which is characteristic of poor nutrition.     

On the timing of moulting processes in reproductively active Northern krill Meganyctiphanes norvegica

The interactions between moult phasing, growth and environmental cues in Northern krill (Meganyctiphanes norvegica) were examined through analysing populations at seasonal, weekly, and daily timescales. The analyses were carried out on resident populations of krill found in three different neritic locations that experience similar environmental signals (the Clyde Sea, Scotland; the Kattegat, Denmark; Gullmarsfjord, Sweden). Seasonal analyses were carried out on the Clyde Sea population and showed that moulting frequency increased significantly moving from winter to summer. The proportion of moulting females in summer samples was often more than double the proportion of moulting males, suggesting that females had a comparatively shorter intermoult period (IMP). Weekly samples taken from the Kattegat showed a similar pattern. However, although the difference between the proportion of female and male moulters was significant in one week, it was not another, mainly because of the variability in the proportion of female moulters. Such variability in females was equally evident in the daily samples taken at Gullmarsfjord. It suggests that females have a shorter IMP (12.5 days) than males (18.4 days) and are more likely to moult in synchrony. Nevertheless, the daily samples revealed that males are also capable of moult synchronisation, although less frequently than females. Shortened IMPs in females were not a result of the abbreviation of specific moult stages. Accordingly, reproductive activity did not alter the course of the normal moult cycle. There was no significant difference between the total body lengths of males and females indicating that females achieve the same levels of growth despite moulting more frequently and having to provision the energy-rich ovaries. This is in contrast to most other crustaceans where the energy costs of reproduction reduce female growth. The fact that females were less abundant than males, probably by suffering a greater level of mortality, suggests that different behavioural strategies, particularly vertical migration regimes, were adopted by each sex to maximise growth and reproduction.     

Moult in relation to some aspects of reproduction and growth in swarms of Antarctic krill,Euphausia superba

In a 14-d period 38 swarms of Antarctic krill,Euphausia superba Dana, were sampled in an area 55.5 x 55.5 km to the southwest of Elephant Island, at the north of the South Shetland Islands. Moult stage, maturity stage, and size of ca. 100 krill from each swarm were measured. Each of the characteristics varied greatly between swarms. Moulting krill were found in most swarms, but in one swarm all of the krill were just about to moult. Ways in which moulting may act as a possible sorting mechanism are discussed. Data for all the analysed krill (ca. 3000 specimens) were used to investigate the interdependence of moult rate, sexual maturation and growth. While all immature krill moulted at approximately the saine rate in the study, there were significant differences in the moulting rates of mature male and female krill. Gravid female krill continued to moult, although less frequently than mature males. As a consequence males had to attach spermatophores to females after each moult. It is likely that variation in moult rate of females and males was related to the energy expenditure required for ovary development in females and spermatophore production and searching behaviour in males. Spawning and moulting were only partly coupled in gravid females. Spawning appeared to take place predominantly during Moult Stage D₂. There was no evidence of intermoult growth by intersegmental dilation.     

Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

Patterns of growth and triacylglycerol content in snow crab Chionoecetes opilio (Brachyura: Majidae) zoeal stages reared in the laboratory

Snow carb Chionoecetes opilio zoea I and zoea II larvae, hatched from females in a controlled mating experiment, were reared in the laboratory at 10.1 °C and 28.0 salinity, to resolve the patterns of growth (dry weight [DW]) and change in energy reserves (triacylglycerols [TAG]) within a given moult cycle. The patterns of growth and change in TAG reserves were similar in each zoeal stage. Following hatching or a moult, the zoeae entered a phase of rapid size increase, i.e. high daily growth rates (5.5 to 12.8% DWd^-1), for 1/3 to 2/5 of the duration of the moult cycle. During the same period, the zoeae accumulated TAG reserves until a maximum (TAG DW^-1) was reached at the end of the phase of rapid growth. The period of high growth rates and of TAG accumulation is interpreted as the required time for the zoeae to reach a point in development [i.e. point of reserves saturation (PRS); Anger and Dawirs (1981)] where sufficient growth and energy reserves allow moulting to the next stage. Following the phase of rapid growth and TAG accumulation, the zoeae entered a phase of low daily growth rates (0 to 1% DWd^-1) during which the TAG reserves decreased to a minimum at the end of the phase. Prior to, and during the moult to zoea II, a phase of negative growth was observed in the zoea I larvae. We conclude that measurement of zoeal size and TAG content, along with morphometric criteria (e.g. epidermal retraction), can be used to assess growth and nutritional condition of C. opilio zoeal stages from the sea.