Population differences in gonad and pyloric caeca cycles of the New Zealand seastarSclerasterias mollis (Echinodermata: Asteroidea)

Inshore and offshore populations ofSclerasterias mollis (Hutton, 1872) were sampled on the outer continental shelf off the Otago Peninsula, New Zealand, and their nutritional and reproductive cycles are described from 1985–1986. Histological changes in the gonads are generally typical of other asteroids. The gametogenic cycle takes 12 mo. The gonad and pyloric caeca indices in both females and males had an inverse relationship.S. mollis accumulates nutrients in the pyloric caeca during summer and early autumn. The gonads develop in the autumn and winter. Offshore seastars were much larger and had significantly higher gonad and pyloric caeca indices than inshore individuals. These differences in body sizes and organ indices arise from differences in food availability and/or population density.     

Changes in the biochemical composition of the pyloric caeca of female seastars,Asterias rubens,during their annual reproductive cycle

The biochemical composition of the pyloric caeca of female seastars (Asterias rubens) was studied throughout the annual reproductive cycle. The pyloric caeca index is high during the pseudo-rest stage of the ovaries and decreases gradually during ovarian growth. The pyloric caeca dry weight varies between 20 and 38% of the fresh weight. Lipids, and less prominently carbohydrates (both glycogen and other reducing sugars), seem to constitute the primary nutrient reserves for gonad growth. Proteins are also available to meet ovarian requirements during vitellogenesis. Glycogen levels are high in late autumn and winter and low at spawning time and in summer, whereas levels of the other reducing sugars are high in post-spawning months and subsequently gradually decrease during gonad development. The total lipid level is extremely variable (averaging 75 mg g^-1 organ fresh weight), peak levels of over 200 mg g^-1 being found in some individuals in summer and at spawning time. The free amino acid level is constant at about 20 mg g^-1. The protein level is rather high throughout the annual cycle, and displays only minor changes. The observed patterns of decrease in major constituents suggest a rapid release of nutrient reserves from the pyloric caeca (deposited during the summer) at the onset of vitellogenesis. Moreover, the needs of the ovaries during further development seem to be so tremendous that during the winter months the digested food may be transported to the ovaries without prior deposition in the pyloric caeca. The changes observed in the pyloric caeca are discussed and compared with those occurring simultaneously in the ovaries.     

Mating behaviour and reproductive cycle of Archaster typicus (Echinodermata: Asteroidea)

This study examines the reproductive cycle, the mechanism of male-on-female pairing behavior and the spawning behavior of Archaster typicus Müller et Troschel. Field studies were conducted in the intertidal zone of the sand beaches at Penghu, Taiwan (23 32N; 119 33E) at ebb-tide in 1984 and 1985. The pairing behavior of A. typicus is a reproductive behavior which leads to simultaneous spawning, increasing the probability of fertilization. As the breeding season approaches, sea stars, especially males, display increased mobility. Because only males tend to mount another individual and because males can detect the sex of another individual by contact with the side of their arms, a male-on-female pair is formed when a male encounters a female. Eighty-five percent of the sea stars observed were paired during the pairing season. Spawning by a paired female is closely followed by spawning of its paired male; male spawning, however, does not induce spawning in its paired female. During spawning, the male turns slightly so that its arms overlap the arched arms of the female. The gonad volume of males is much less than that of females. This may result from the high efficiency of fertilization in this species, which does not require a large amount of gametes to be released, or from the higher energetic demands made on the males.     

Seasonal changes in pyloric caecum and gonad indices during the annual reproductive cycle in the seastar Asterias forbesi

Seasonal changes in pyloric caecum and gonad indices were studied in a population of the seastar Asterias forbesi (Desor) from East Rockaway Inlet (Long Island, New York) during two annual cycles (1980/81, 1982/83). Pyloric caeca indices increased during fall, reached a maximum about April, and declined sharply to a minimum in mid-summer. Gonad indices increased during fall and winter and reached a maximum about May. Judging from gonad size analysis, spawning occurred in late June to early July, at bottom temperatures of 16° to 18°C. There was no long-term inverse relationship between pyloric caecum and gonad indices. Since nutrients and energy ingested during the important fall feeding period are utilized simultaneously for body growth and gonad development, it is unnecessary for the pyloric caeca to store nutrients over long periods as occurs in A. rubens and many other species. The specific caeca-gonad relationship is probably an adaptation to the extreme seasonal thermal fluctuations of the NW Atlantic, which constrain the feeding activity of A. forbesi and limit the possibility of long-term storage. In this population, most individuals grow rapidly in their first year (including first two summers) and spawn in their third summer. Few seastars survive to spawn again in their fourth summer. However, gametogenesis may take place in some individuals in their second summer. Year-to-year variability in mean size at spawning may reflect temporal variation in environmental conditions (weather, food availability) at this site.     

Storage and depletion of lipid components in the pyloric caeca and ovaries of the seastarAsterias rubens during its annual reproductive cycle

Detailed information is presented on the changes in lipid-class profiles (polar lipids, total neutral lipids, free sterols, steryl esters, triacylglycerols, neutral ether lipids and free fatty acids), which are manifest during the annual reproductive cycle of the seastarAsterias rubens for both the storage organs (pyloric caeca) and utilizing organs (ovaries). The ovaries appear to accumulate all individual lipid components proportional to progression of vitellogenesis. Pyloric caeca, in contrast, show variable lipid-class levels, probably due to more complex processes in these organs, operating simultaneously: digestion of food, accumulation of storage lipids, mobilization of stored lipids and their release. The results are discussed in terms of current knowledge on the role of lipids in echinoderm reproduction.     

The annual reproductive cycle of the holothurian Aslia lefevrei (Dendrochirota: Echinodermata)

The dendrochirote holothurian Aslia lefevrei (Barrois) is locally and densely aggregated (>70 individuals m^-2) in Galway Bay, on the west coast of Ireland. It has a discrete, relatively short breeding period from the end of February to the beginning of April, with peak activity in March. The sexes are separate and the single gonad, consisting of numerous tubules (100 to 800), lies in the anterior coelom. Both gonad index values and histological observations indicate that gametogenesis resumes in June/July after a brief post-spawned resting period. Little oocyte growth and sperm production occurs over the warm summer months when the holothurian is actively feeding. Gonad index values, however, increase rapidly during this period and reach high values by early winter without concomitant increases in maturity indices. This is subsequently followed by slight reductions in the gonad indices, before reaching peak values in February. It is suggested that this is probably due to the initial build-up and subsequent assimilation of storage products in the gonad, the development of gametogenic products and the frequent assimilation thereafter of unwanted ova. Spawning generally takes between 5 and 60 min, with the entire contents of the gonad rarely being shed in a single release. During this process, the holothurian raises its anterior end and, with the extended tentacles close together, emits a stream of gametes from the genital pore in the mid-dorsal line. Laboratory experiments and general observations indicate that increased temperature and low light levels may influence spawning behaviour.Contribution No. 248 to the School of Marine Sciences, University College Galway, Ireland     

Reproductive cycle of the brooding echinoid Abatus cordatus (Echinodermata) in Kerguelen (Antarctic Ocean): changes in the organ indices,biochemical composition and caloric content of the gonads

Variations in organ indices, organic levels, organic and caloric contents of the gonads of the brooding echinoid Abatus cordatus (Verrill), collected in the Morbihan Gulf (49°30′S; 70°E) were studied over a 1 yr period (March 1978 to January 1979). In both sexes, a clear annual cycle was discernable for organ indices, and for organic and caloric contents. In contrast, the protein levels in the gonads varied moderately, while the levels of lipid and carbohydrate remained almost constant. This peculiarity sets A. cordatus apart from nearly all other echinoids characterized by an annual reproductive cycle. The male and female gonad indices change synchronously. The gonad indices double during maturation, reflecting variations in the germinal tissue. In both sexes, the maximal gonad index (at the end of March) coincides with the onset of the spawning season. The average male gonad index is 50% higher than that of the female. This difference is related to the restriction of coelomic space in the female due to development of the brood pouches. At maturity, the lipid and carbohydrate levels in the ovary are 2.8 and 2.1 times greater, respectively, than the levels in the testis, the latter having a 25% higher protein level. This disparity in organic levels reflects mainly the differences in germinal tissue composition. The female reproductive effort is higher than that of the male. The spawn energy output of a standard individual is 200 cal for a male and 228 cal for a female. The unusually lower gonad index of the female is compensated for by higher lipid and carbohydrate levels, giving the ovary a 40% higher calorific value.     

The reproductive cycle of Asterina stellifera (Möbius) (Echinodermata: Asteroidea) in the Cabo Frio region, southeastern Brazil

Reproduction of Asterina stellifera was studied in two populations from the Cabo Frio region, southeastern Brazil. One was located in an intertidal seagrass flat at Japonês Island (1994–1995); the second was on a sublittoral rocky shore at Pontal beaches (1999–2000). Reproductive cycle was studied at monthly intervals by organ indices and histological examinations of gonads. Although these two populations were studied in different habitats and at different times, the periodicity of the annual reproductive cycle was very similar for the two populations. The highest gonad index (GI) values were found in August/September (late winter), followed by a drop in GI values, indicating spawning. The GI and pyloric caeca index (PCI) tended to have an inverse relationship in both populations, but it was only statistically significant in the rocky-shore population. Gametogenesis in the two populations was described and roughly confirmed the reproductive cycles determined by GIs. Published online: 9 August 2002     

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.     

Changes in the biochemical composition of the ovaries of the seastar Asterias rubens during its annual reproductive cycle

The biochemical composition of the ovaries of the asteroid echinoderm Asterias rubens was studied, with emphasis on the changes which are manifest during the annual reproductive cycle. Ovarian dry weight displays only minor changes during ovarian development, constituting about 20% of the organ fresh weight. Levels of all constituents show some variation, but in general increase with ovarian growth. Only the free amino acid level is rather constant. Except for glycogen, the contents of other constituents (total lipids, other reducing sugars, free amino acids and proteins) show a rectilinear increase with ovarian growth. Glycogen content increases rectilinearly with time during early vitellogenesis, but remains constant during further development until ovarian maturity. By plotting constituent levels and contents against a non-linear gonad index axis rather than against sampling date, results can be correlated more easily with ovarian growth stages. The possibility of endocytosis of a vitellogenin-like yolk precursor by the ovary is discussed.     

Annual reproductive cycle, spatial distribution, abundance, and size structure of Oreaster reticulatus (Echinodermata: Asteroidea) in Bocas del Toro, Panama

The structure, distribution, and population abundance of Oreaster reticulatus (Linnaeus, 1758) in 47,157 ha of shallow-water habitat in the archipelago of Bocas del Toro, Panama, were assessed from May to October 2000. The reproductive cycle of the sea star was studied in Isla Solarte, from February 2000 to February 2001. In total, 4,818 sea stars were recorded with a mean density of 149.7 ind. ha^-1, and a population of over 7 million was estimated for the archipelago. O. reticulatus was absent in ca. 50% of the evaluated areas, possibly due to high runoff and sedimentation; highest density was observed in an intermediate-runoff regime (255 ind. ha^-1). About 45% of the population was found in substrata dominated by seagrass (Thalassia testudinum) and coarse, calcareous sand, 51% occurred in habitats where coral reef patches were mixed with seagrass, and 4% exclusively on coral reefs. The average size, based on the major radius, was 9.5 cm (3-21 cm), with a population structure composed of ca. 83% juveniles and 17% adults. The average reproductive size, measured as major radius, was 15 cm, and the minimum was 7 cm. Both males and females with a maximum stage (IV) of gonad development were observed throughout the year. The gonad index showed three peaks of maximum reproductive activity, which is not comparable to studies from other localities. The reproductive cycle did not seem to be related to water temperature, which ranged from 27°C to 30°C, but may respond more closely to changes in local rainfall. This relationship was not statistically significant based on this 1-year study. These data provide a useful baseline for management of local populations in the face of an increasing harvest for the aquarium trade and as souvenirs.     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Reproduction and annual gametogenic cycle in Nephtys hombergii and N. caeca (Polychaeta: Nephtyidae)

The gametogenic cycles of Nephtys hombergii Audouin et M. Edw. and N. caeca Fabricius are described. Both species are polytelic (=discrete, iteroparous) and breed in their second and subsequent years. They spawn in May in the Tyne estuary. After spawning, N. hombergii retains a large number of gametocytes, which are ultimately resorbed. A new generation of gametocytes is proliferated by September and mature gametes accumulate during the winter months. Gametic development is not well synchronised.     

Abundance,spatial distribution,and size structure of populations of Oreaster reticulatus (Echinodermata: Asteroidea) on sand bottoms

Three populations of Oreaster reticulatus (Linnaeus, 1758) inhabiting shallow-water (<4 m) seagrass habitats in the Grenadines (West Indies, Caribbean Sea) were associated predominantly with beds of Halodule wrightii. Occupation of fringing inshore areas of bare sand was inversely related to wave action; even where sandy patches occurred offshore, the preferred substratum was H. wrightii. The association of O. reticulatus with H. wrightii is related to the asteroid's microphagous feeding habit and the availability of food resources associated with the seagrass. O. reticulatus rarely occurred on dense beds of Thalassia testudinum, but was moderately abundant in areas of sparse cover. Differences in the occurrence of O. reticulatus among seagrass types may be related to factors afdecting foraging effort, such as the tractability of the substratum and mobility upon it. Populations of O. reticulatus exhibited an aggregated dispersion within beds of H. wrightii, possibly attributable to local substratum heterogeneity and/or reproductive behavior. Increased turbulence induced migration to deeper water and markedly increased aggregation along offshore boundaries. The populations were primarily adults, with some late juvenile stages. The paucity of juveniles and their cryptic behavior and coloration suggest that settlement and early postmetamorphic development occurs in alternate habitats, such as dense beds of T. testudinum. Interpopulation differences in size structure may be associated with differences in the quality and availability of food sources.     

Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico

Two morphs of the asteroid genus Echinaster, E. Type 1 and E. Type 2, have reciprocal annual cycles of reproduction and nutrient storage. The pyloric caeca (storage organs) reach maximal size over winter then decrease with increasing gametogenic activity in the spring. E. Type 1 broadcasts buoyant eggs in late May to early June. E. Type 2 deposits benthic eggs in late April. Lecithotrophic larval development is similar in both morphs except that larvae of E. Type 1 undergo a brief planktonic phase. E. Type 1 reaches sexual maturity at a larger size, expends a lower reproductive effort but has a greater absolute reproductive output than E. Type 2. E. Type 2 produces fewer, larger eggs and has a greater parental investment per egg. In both morphs, females have much higher gonadal lipid levels than males and expend a higher reproductive effort in terms organic matter and energy. Reproductive effort, reproductive output, nutrient storage in the pyloric caeca and body size varied between populations and between years in each morph. Egg size and parental investment per egg were constant. Within-morph variability is attributed to differences in nutritional state. Differences in reproductive strategy support the hypothesis that the morphs are separate species.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Seasonal changes in the biochemical composition of the chiton Chiton iatricus (Polyplacophora: Mollusca) and the marine pulmonate Onchidium verruculatum (Gastropoda: Mollusca) in relation to their reproductive cycles

Seasonal changes in the biochemical composition of two molluses, gonochoric Chiton iatricus and a marine pulmonate, Ochnidium verruculatum inhabiting the same tropical area, but having different modes of reproduction, were studied. Protein is the major biochemical component stored in C. iatricus, whereas lipid is predominantly stored in O. verruculatum. The correlation between these quantifications and the annual reproductive cycles of the two species investigated suggest that nutrients stored in the somatic body parts are channeled towards the gonad during the gametogenesis. In C. iatricus sexual differences efrist in the biochemical needs of the ovary and the testis. The ovary always has a higher protein content than the testis. In the hermaphrodite pulmonate O. verruculatum, biochemical expenditure on gamete production is relatively low because this species has to spend energy on the reproductive propagation viz. copulation and egg-laying. On the contrary, in C. iatricus the biochemical energy is mainly utilized for the production of gametes which have to be produced in great numbers in order to counter-balance the loss of gametes caused by external fertilization. The collected data are discussed in the light of comparative physiology of reproduction.     

Costs of extended brood protection in the Antarctic sea star, Neosmilaster georgianus (Echinodermata: Asteroidea)

Changes in size, morphology, and biochemical composition in adults and embryos of a brooding sea star, Neosmilaster georgianus (Sladen), were studied in a population adjacent to Anvers Island, Antarctic Peninsula (64°46′S; 64°04′W) during the austral spring, 1991. Five morphological stages of development were designated in 24 broods, and for each the weight and biochemical composition of the brooding adults and their embryos were determined. Between Stage 1 and 2, the dry weight (dw) and organic weight (ow) of the embryo did not change. From Stage 2 to 3, the dw and ow increased significantly by 10%. Stage 2 and 3 embryos were in clusters of a few (2 to 10) to as many as 40 individuals. In the smaller clusters, individual embryos were attached by tissue cords to another, sometimes atrophied, brood member. In the larger clusters, they were attached to a central mass of tissue containing remnants of embryos. We interpret these interactions as a form of cannibalism which may account for the weight gains between Stage 2 and 3. During Stages 4 and 5, as juvenile form was approached, the dw and ash weight of the young increased significantly and the ow decreased significantly. The calculated energy content for the juvenile (Stage 5) was not significantly different from the energy content of the earliest undifferentiated stage (Stage 1), an indication that most of the organic matter in the egg is the primary contribution to the large juvenile. In brooding females, pyloric caeca indices declined by 52% from Stage 1 to Stage 5 and pyloric energy stores declined by 63% due to proportionately equivalent declines in protein and lipid. The ovary index was low and increased only slightly during brood protection, while the size of the largest oocytes remained approximately 23% that of ova. Energy stores in the pyloric caeca of brooding N. georgianus thus become depleted over a long period of incubation during which the adults apparently starve. This may delay oocyte development and ultimately limit the frequency of individual reproduction. Received: 15 September 1998 / Accepted: 9 March 1999     

Seasonal variations in condition,reproductive activity and biochemical composition of the flat oyster,Ostrea edulis,from San Cibran (Galicia,Spain)

The influence of environmental parameters (temperature, salinity and available food) on the condition, reproductive activity and biochemical composition of a native population of Ostrea edulis L. in San Cibran (Galicia, Spain) was studied between September 1988 and December 1989. Histological preparations of gonads showed that gametogenesis started when temperature was at its lowest in winter. The water temperature in San Cibran never fell low enough to interrupt it. Gametogenesis proceeded slowly and spawning took place in May–June, although the predicted time of ripening was early March. Salinity in San Cibran was relatively stable throughout the year; it did not seem to have any influence on gametogenesis. Available food appeared to be a very important factor in controlling gonad growth, once gametogenesis was initiated. The major concentration of suspended organic particulate matter was present in the spring at the time of rapid gonadal maturation. Only one spawning period was observed. Larvae were released when the food in the water was high. Seasonal changes in the main biochemical components of this oyster were determined for a standard individual. Lipids and carbohydrates presented a similar time-course whereas proteins were constant. When food was abundant, energy reserves were built up. Spawning produced a decrease in biochemical constituent levels, and recovery coincided with the phytoplankton bloom. The stored reserves, mainly lipids, were used to overcome a state of energy imbalance in late autumn associated with low food availability. Results show this oyster to be an opportunist organism which concentrates its reproductive effort during a short period of favourable conditions and which is directly dependent on nutritive availability in the environment.     

Interannual variation in the reproductive cycle and biochemical composition of the cockle Cerastoderma edule from Mundaca Estuary (Biscay,North Spain)

Seasonal determinants of body weight, biochemical composition and reproductive condition in the cockle Cerastoderma edule L. from the Mundaca Estuary, Spain, were performed from December 1983 to July 1985. Interannual differences in timing of gametogenesis and spawning between 1984 and 1985 were correlated with temporal variations in the cycles of storage and utilisation of carbohydrate reserves. Young individuals exhibited larger annual fluctuations in soft-body weight, displaying higher growth rates in spring and greater diminishment of energy reserves during autumn and winter than older individuals. Carbohydrate content displayed a similar tendency, levels being lower in young individuals throughout the year.