Glycogen storage in relation to the moult cycle in the two crustaceansEmerita asiatica andLigia exotica

InEmerita asiatica, the quantitative fluctuations in the glycogen content of the hepatopancreas were markedly related to the moult cycle. The glycogen content of the hepatopancreas was maximum during premoult stages. It has been suggested that reserve glycogen, in addition to meeting general metabolic needs, may be especially involved in the genesis of the sugar precursors of chitin. The marked fall in the hepatopancreas glycogen values following moulting is probably due to its utilization in chitin synthesis during and immediately following moulting. InE. asiatica, the storage of glycogen was found to be meagre in muscles, and muscle glycogen had no bearing on chitin synthesis during moulting. InLigia exotica, glycogen is stored to a lesser extent than inE. asiatica, and was found in connective tissue and muscles. Significant fluctuations were noted in glycogen values correlated with the phases in the moult cycle when chitin synthesis took place. InL. exotica, the hepatopancreas does not serve as a storage depot for glycogen related to chitin synthesis.     

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.     

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.     

Effect of the insect growth regulator diflubenzuron (Dimilin®) on the uptake of glucose and N-acetylglucosamine into the cuticle of crab larvae

Incorporation of ³H-labelled glucose and ³H-labelled N-acetylglucosamine (NAGA) — both precursors to chitin —into the cuticle of Rhithropanopeus harrisii (Gould) larvae (Crustacea: Brachyura) has been examined at different stages of the moult cycle in control larvae as well as in larvae treated with the insect growth regulator diflubenzuron (Dimilin^®). As far as the control larvae were concerned, the incorporation of both precursors was high at the postmoult stage when endocuticle was secreted. NAGA appeared to be a more specific precursor of cuticular material than glucose during the premoult stage when exocuticle was produced. Incorporation of both precursors was low immediately before ecdysis and during the intermoult stage when secretion of the cuticle is complete. The results show that incorporation of glucose into chitin was greatly inhibited by the pollutant during the postmoult stage when endocuticle is produced, while incorporation of NAGA was reduced to a lesser extent at this stage. Diflubenzuron treatment markedly affected the incorporation of both NAGA and glucose in the premoult stage during secretion of exocuticle.     

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     

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.     

Nucleic acids and growth of larval and early juvenile spider crab,Hyas araneus

The spider crabHyas araneus (L.) was collected from the North Sea in winter 1986–1987 and reared in the laboratory from hatching of the Zoea I (ZI) through the first juvenile instar (CI). Within a given moult cycle, individuals of the same age were sampled in intervals of 2 (ZI, ZII, CI) or 3 d (megalopa) for analysis of dry weight (W), carbon (C), nitrogen (N), hydrogen (H), protein, DNA, and RNA. Lipid was calculated from C. Biomass, growth rate and nucleic acid contents showed high variability during each moult cycle and between instars. Instantaneous growth rates of C were high in postmoult and intermoult, and low in the premoult period of each moult cycle. A shift was observed from high rates of lipid accumulation in the postmoult and intermoult stages to proportionally increasing protein accumulation during late premoult (ZI), or throughout a major part of the remaining moult cycle (in all other instars). DNA was accumulated throughout the ZI and ZII instars, but decreased in late premoult megalopa. It increased again from late intermoult through intermediate premoult in juveniles. RNA increased continuously during ZI and ZII, and decreased in the megalopa, almost to levels that had been found immediately after hatching. In juveniles, variation in RNA followed closely those in DNA. Cell multiplication (expressed by DNA increase) dominated over increase in cell size (defined by the C/DNA ratio) during the zoeal instars and in postmoult through early intermoult in the megalopa and CI. When specific (C-related) RNA values and RNA/DNA ratios were compared with instantaneous growth rates in C and N, no general correspondence was detected. The only significant relationship between specific RNA values and instantaneous C or N growth rates was found in the megalopa. The same held for the relationship between the RNA/DNA ratio and growth. Here, in addition to the megalopa, a correspondence with C growth was also found in the CI instar. Our results suggest that variation in nucleic acids may provide useful insights into mechanisms of growth on the cellular level (cell multiplication vs cell enlargement). However, lack of general correlation with variation in growth rates ofH. araneus larvae shows that the use of nucleic acids as a measure of growth is probably based upon too simplistic assumptions; it may not yield reliable predictions, when growth is associated with developmental events.     

The essential amino-acid requirements of the prawnPalaemon serratus. The growth of prawns on diets containing proteins of different amino-acid compositions

Two specimens ofPalaemon serratus Pennant (of about 4 g wet weight) were each given intrahaemocoelic injections of [U-¹⁴C] acetate. They were maintained under controlled conditions, and the release of¹⁴CO₂ and other labelled metabolites was followed for 6 days. The prawns were then sacrificed and fractionated into trichloroacetic acid soluble compounds, lipids, nucleic acids and a protein/chitin residue. Little or no¹⁴C had been incorporated into arginine, methionine, valine, threonine, isoleucine, leucine, lysine, histidine, phenylalanine and tryptophan. It was inferred thatP. serratus cannot synthesize these amino acids from ordinarily available materials, and that the prawns have an absolute dietary requirement for them. The glucosamine isolated from the protein/chitin residue contained more radioactivity than any of the other compounds examined. This implies a rapid turnover of glucosamine and chitin inP. serratus. The gain in weight of prawns fed on compound diets containing freeze-dried cod muscle was about 70% that of prawns fed on fresh mussel mantle. Predigestion of freeze-dried cod muscle with proteolytic enzymes did not significantly enhance its nutritional valve as assessed by increases in weight. Prawns fed two other compounded diets containing proteins of low nutritional value (gelatin and zein, respectively) grew at about 1/5 the rate of prawns fed fresh mussel mantle. Supplementation of the gelatin and zein in these diets with tryptophan, and with lysine plus tryptophan, respectively, failed to augment their nutritional value significantly.     

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.     

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.     

Copper and zinc handling during the moult cycle of male and female shore crabs Carcinus maenas

Tissue concentrations and contents of copper and zinc were determined at 11 different stages of the moult cycle in male and female shore crabs Carcinus maenas. Metal concentrations in haemolymph, gills, midgut gland, muscle, and exoskeleton of males and haemolymph, gills, and midgut gland of females were determined, as were haemocyanin concentrations and haemolymph volumes (using ¹⁴C-inulin) in males. The changes in tissue Cu and Zn concentrations and contents that occur throughout the moult cycle can be attributed to muscle breakdown in late premoult, the period of starvation in late premoult and early postmoult, the resorption from and shedding of the old exoskeleton, and the dilution of the haemolymph caused by water uptake around the time of ecdysis. The present study demonstrates that whole-body Cu and Zn contents remain constant during a large part of the moult cycle of male and female C. maenas. This state of whole-body trace metal homeostasis is maintained in spite of major changes in tissue proportions and tissue Cu and Zn concentrations and contents. Previous studies have not carried out the necessary analysis to move from theoretical estimates to quantitative determination of the changes in tissue metal distribution associated with moulting in crustaceans; the data presented illustrate the necessity of measuring both tissue concentrations and contents of metals to avoid misinterpretation of either.     

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.     

Setal development,moult-staging and ecdysis in the banana prawn Penaeus merguiensis

Specimens of juvenile Penaeus merguiensis de Man (1888) were captured by seine netting from the shores of Cleveland Bay, Townsville, Queensland Australia between February 1980 and November 1981. Accurate determination of moult cycle stage can be achieved rapidly by microsopical examination of setal development in an excised whole pleopod. Four major stages and 5 substages within premoult were clearly defined. The rate of setogenesis varies along the length of a pleopod, precluding the examination of only one specific location on the pleopod for accurate moult cycle staging. The process of ecdysis lasts for no more than 40 s.     

Ultrastructure of the integument of a pelagic Crustacean: moult cycle related studies on the Antarctic krill,Euphausia superba

The ultrastructure of euphausiid integument was examined in relation to the moult cycle and supplemented by investigations of chitinase activity in the integument and content of N-acetyl--D-glucosamine in the hemolymph. The Antarctic krill, Euphausia superba was collected in 1983 in Admiralty Bay, King George Island, Antarctica. Some specimens of the Northern krill, Meganyctiphanes norvegica, from the Danish Kattegat served for comparison. As a major aim of the study, the moult staging system developed for living tissue could be verified by ultrastructural findings. Under experimental high production conditions of the Antarctic summer, no period of rest or intermoult between post- and premoult was observed in subadult E. superba. Neither was a resting phase seen at the cellular level, the epidermis remained active. The epidermal gland cells did not show any cyclical changes, and the organelles of protein synthesis were generally well developed in all moult stages. In order to follow the physiological course of events, structural and biochemical methods were combined and showed as a result that the last moult stage before ecdysis is characterized by massive cuticular resorption. The epicuticle remained ultrastructurally unchanged before and after ecdysis, even though its permeability should alter at ecdysis. The existence of muscle insertions which connect the old and the new cuticle across the exuvial space suggests an answer to the question why E. superba is hardly impaired in swimming almost up to the time of ecdysis.Dedicated to Professor Dr. H. Leonhardt on the occasion of his 70th birthday     

Light quality in relation to growth,photosynthetic rates and carbon metabolism in two species of marine plankton algae

The effect of light quality on growth, photosynthesis and carbon metabolism in two species of marine algae,Cyclotella nana (Hustedt) andDunaliella tertiolecta (Butcher), was examined. Relative growth constants forC. nana were 0.37, 0.29 and 0.25 in blue, white and green light, respectively. Corresponding constants were 0.41, 0.31 and 0.29 forD. tertiolecta. Photosynthetic rates in both species were higher in blue light and lower in green light compared with white light of the same intensity. More than 60% of¹⁴C assimilated byC. nana orD. tertiolecta grown in blue or green light was incorporated into the ethanol-insoluble fraction, compared with 10 to 30% in this fraction in white light. The relative importance of the various components within this fraction was independent of light quality. Although less¹⁴C was assimilated into the ethanol-soluble fraction in blue or green light, there was a relative increase in some amino acids and organic acids in this fraction and a decrease in sugars and sugar phosphates relative to white light of the same intensity. These differences were independent of light intensity, photosynthetic rate and cell density in the cultures.     

Glycohistochemistry of a marine sponge, Chondrilla nucula (Porifera, Desmospongiae), with remarks on a possibly related antimicrobial defense strategy and a note on exopinacoderm function

Based on carbohydrate histochemistry, including the use of lectins, and TEM, the study describes the distribution of terminal sugars in different structures of the demosponge Chondrilla nucula. The results of the general and specific carbohydrate histochemical approaches confirmed the presence of acidic and neutral glycoconjugates in the cells, and, with declining amounts from the ectosome to the mesohyl, in the extracellular matrix (ECM). AB-PAS staining indicated acidic complex carbohydrates particularly in the exopinacoderm, and more neutral ones in the cells and the ECM of the mesohyl. The PO-lectins applied demonstrated a general spectrum of free saccharide residues (α-d-mannose, α-/β-d-N-acetylglucosamine, α-d-N-acetylgalactosamine, α-d-galactose, β-d-galactose) in both sponge parts; α-l-fucose was only distinct in the ectosome. Sialic acids [siaα(2,3)-galactose, siaα(2,6)-N-acetylgalactosamine] were dominant in the very thin exopinacoderm, indicating O-linked high molecular weight glycoproteins. In this way a glycophysiologically ‘rigid’ outer mucus cover is developed as protection against mechanical hazards. Some of the free sugars (α-d-mannose, N-acetylglucosamine, N-acetylgalactosamine β-d-galactose, α-l-fucose) are known to prevent the adherence of different bacteria and fungi to cellular surfaces. Thus a high concentration of such sugars, may impede massive attacks of micro-inhabitants on mobile sponge cells, pinacocytes, and the exopinacoderm layer.     

Studies on the transport of sugars in the holothurian Holothuria scabra

Experiments using ¹⁴C sugars were carried out on the holothurian Holothuria scabra Jäger, in order to assess the role of its perivisceral fluid in the translocation and transportation of dissolved organic materials. The results obtained indicate that the perivisceral fluid plays a significant part. Rates of ¹⁴C glucose, fructose and sucrose uptake were followed in major tissues such as the alimentary canal, haemal system, respiratory tree and body wall. The monosaccharides are absorbed more intensively by digestive and haemal systems, the disaccharide by respiratory tree and body wall. It is, therefore, presumed that tissues in H. scabra selectively absorb sugars depending on their metabolic activity.     

The histology,histochemistry and ultrastructure of the cement apparatus of three adult sessile barnacles,Elminius modestus,Balanus balanoides and Balanus hameri

The histology, histochemistry and ultrastructure of the cement gland cells of 3 species of sessile barnacles are described. Cells of both Elminius modestus Darwin and Balanus balanoides (L.) (membranous bases) are similar histologically and ultrastructurally, and within the cells there are 2 distinct regions. One is the site of secretion accumulation, the other the site of secretion synthesis. In Balanus hameri Ascanius (calcareous base) the cells are different in that the cytoplasm is more homogeneous. In all 3 species protein is the final secretion product (cement) of the cement cells. The mode of discharge of secretion from the cells differs between the membrane based and calcareous based barnacles. In the former, there is a series of collecting canals within the cytoplasm of the cement cells (intracellular canals) which join with the larger extracellular cement ducts. Secretion passes into the intracellular canals and is moved along to the larger cement ducts, which have an inner chitin lining. In Balanus hameri (calcareous base) there are no intracellular collecting canals. Secretions are thought to pass from the cement cells into the cement duct cells direct. The cement ducts in this species are not chitin lined.     

A new method to determine in situ growth rates of decapod shrimp: a case study with brown shrimp Crangon crangon

Shrimps are economically and ecologically very important, yet a lack of ageing techniques and hence unknown growth rates often impairs analytical assessments and management. A new method for the determination of in situ growth rates of shrimps is presented, based on dry weight condition. Since this index oscillates from low values directly after moult to highest values prior to moult in constantly feeding shrimp, the lowest observed pre-moult condition followed by a moult was introduced as a reference value to separate growing and starving individuals in field data. Experiments with Crangon crangon confirmed that (1) post-moult condition varies in a narrow physiologically optimal range, regardless of recent growth increments, and (2) dry weight condition prior to moult is closely related to the subsequent length increment. The method was applied to estimate growth increments from in situ dry weight condition data of C. crangon. The new method can easily be applied to other related species, since the required data can be obtained from very simple short-term experiments.     

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