Patrick Mayzaud

Lipids and life strategy of Arctic Calanus

Abstract The three Arctic Calanus species, C. finmarchicus (Gunnerus, 1765), C. glacialis (Jaschov, 1955), and C. hyperboreus, are the most important herbivores in Arctic seas in terms of species biomass. They play a key role in the lipid-based energy flux in the Arctic, converting low-energy carbohydrates and proteins in ice algae and phytoplankton into high-energy wax esters. In this paper we review the over-wintering strategy, seasonal migration, stage development, life span, feeding strategy, body size, lipid biochemistry and the geographic distribution of the three dominant Calanus species in Arctic waters. We then relate these parameters to other biotic and abiotic factors, such as the timing of the Arctic phytoplankton and ice algae bloom, sea ice cover and climate variability. We also present new data on fatty acid and fatty alcohol content in the three Calanus species in addition to reviewing the available literature on these topics. These data are analysed for species homogeneity and geographic grouping. The dominance of diatom fatty acid trophic markers in the lipids of Calanus underpins the importance of diatoms as Arctic primary producers, even if dinoflagellates and Phaeocystis pouchetii can also be important food sources for the calanoid copepods. We conclude that the Arctic Calanus species are herbivores, engineered to feed on the Arctic bloom, and that the timing of the bloom is the most important factor in determining the life strategies of the individual species.

Respiration and nitrogen excretion of zooplankton. IV. The influence of starvation on the metabolism and the biochemical composition of some species

Changes in the respiration, ammonia excretion and biochemical composition were studied for three species of starving zooplankton (Calanus finmarchicus, Sagitta elegans, and Acartia clausi). Over the period of starvation, the respiration rate of all three species followed the same pattern of an initial decrease followed by a more or less constant level. A similar pattern was observed for the ammonia excretion rate of S. elegans and A. clausi, whereas C. finmarchicus excretion appeared to oscillate between high and low levels of protein catabolism. Study of the biochemical changes showed that C. finmarchicus consumed primarily lipids, and at times proteins, to meet its energy requirement whereas S. elegans and A. clausi primarily used protein. Variations in the elemental composition as well as the O:N ratio confirmed that C. finmarchicus alternated between periods of protein-dominant catabolism and lipid-dominant catabolism during starvation. No similar change in catabolism was observed in the two other species. The results are discussed in terms of physiological mechanisms of resistance to starvation and were used to calculate the energy budget of S. elegans and C. finmarchicus during the period of total starvation. The significance of such budgets is discussed and some of the sources of error examined.

some aspects of the biochemical and mineral composition of marine plankton

Abstract A natural population of phytoplankton and two species of zooplankton, Calanus finmarchicus (Gunnerus) (stage V) and Sagitta elegans (Verrill) (mature), were analysed for their biochemical and mineral composition. Comparison with published data for the same categories of plankton shows that the distribution of both organic and mineral fractions is highly variable, the causes of which are not equally understood. The composition of the organic fraction is affected by the duality of function of the biochemical constituents and by the large number of ecological factors which influence their concentration. Whether this applies to the mineral fraction remains uncertain; nevertheless it appears that plankton concentrate trace metals, that phytoplankton has a greater ability to concentrate them than zooplankton, and that the ability of zooplankton to concentrate various metals varies with the species.

Composition and community structure of pelagic copepods in the Indian sector of the Antarctic Ocean during the end of the austral summer

The present paper describes latitudinal and vertical changes in the composition, abundance and diversity of copepods in the Indian sector of the Antarctic Ocean, during the end of austral summer along a transect on 66°30′E between 43 and 62°S, within three layers (600–0, 200–0, 100–0 m). Highest copepod densities were noted in the central part of the transect, between the Antarctic Divergence and the Antarctic Convergence, with a maximum in the Antarctic Divergence zone, particularly in the upper levels of the water column. A total number of 80 copepod species were identified over the entire survey area. The south end and the central part of the transect comprised a small number of species. North of the Antarctic Convergence, this number increased markedly with the progressive disappearence of those species characteristic of Antarctic waters and their replacement by temperate and subtropical species. Generally, small copepods, particularly Oithona similis, Oithona frigida and Ctenocalanus citer, dominated in numbers in both Antarctic and sub-Antarctic areas. The contribution of large species to total copepod numbers was much lower, with Calanus simillimus in the central part of the transect, Pleuromamma borealis in the subtropical zone and Calanus propinquus in the southern part. Correspondence analysis showed a marked latitudinal gradient in population structure with four groups of samples and species corresponding to four latitudinal zones. Community structure (species richness, relative dominance index, evenness, Shannon species diversity index) and species abundance patterns (as rank-frequency diagrams) suggested that the maturity and species richness increased gradually from south to north. A low diversity index and evenness were observed in the area of the Antarctic Divergence, whereas the convergence zone showed high diversity and evenness. Conversely, the frontal zone showed high diversity and evenness. Distribution appeared unrelated to chlorophyll concentrations and on the large scale was related to the hydrologic characteristics.

DIEL VARIATIONS OF CARBOHYDRATES AND NEUTRAL LIPIDS IN NITROGEN-SUFFICIENT AND NITROGEN-STARVED CYCLOSTAT CULTURES OF ISOCHRYSIS SP.(1).

The goal of this study was to investigate the time response of two major carbon (C) reserves, respectively neutral lipids (NL) and total carbohydrate (TC), in the Haptophyte Isochrysis sp. growing in nitrogen (N)‐sufficient or N‐starved conditions and under light:dark (L:D) cycles. Experiments were carried out in a cyclostat culture system that allowed the following of the dynamics of the main cell compounds at both hourly and daily time scales. Under N‐sufficient conditions, the L:D cycles cause the population to be synchronized, with most of the cells dividing at the beginning of the dark period. The C‐specific growth rate was maximal around midday and negative during the dark period due to respiration processes. NL and TC both accumulated during the day and consumed during the night. We showed that NL and TC are highly dynamic compounds, as more than three quarters of NL and TC accumulated during the light period were consumed during the dark period. In contrast to NL, phospholipid and glycolipid to C ratios remained quite stable during the light/dark cycles. The major effect of N starvation on the NL and TC dynamics was to uncouple their diel variations from the L:D cycle, in two different ways depending on their respective role during short‐term acclimation. Whereas the TC per cell ratio increased rapidly to reach a stable value in response to N starvation, NL per cell continued to oscillate, but with a pattern out of phase with the L:D cycle.

NEUTRAL LIPID AND CARBOHYDRATE PRODUCTIVITIES AS A RESPONSE TO NITROGEN STATUS IN ISOCHRYSIS SP. (T-ISO; HAPTOPHYCEAE): STARVATION VERSUS LIMITATION(1).

Partitioning of the carbon (C) fixed during photosynthesis between neutral lipids (NL) and carbohydrates was investigated in Isochrysis sp. (Haptophyceae) in relation to its nitrogen (N) status. Using batch and nitrate‐limited continuous cultures, we studied the response of these energy reserve pools to both conditions of N starvation and limitation. During N starvation, NL and carbohydrate quotas increased but their specific growth rates (specific rates of variation, μCAR and μNL) decreased. When cells were successively deprived and then resupplied with NO3, both carbohydrates and neutral lipids were inversely related to the N quota (N:C). These negative relationships were not identical during N impoverishment and replenishment, indicating a hysteresis phenomenon between N and C reserve mobilizations. Cells acclimated to increasing degrees of N limitation in steady‐state chemostat cultures showed decreasing NL quota and increasing carbohydrate quota. N starvation led to a visible but only transient increase of NL productivity. In continuous cultures, the highest NL productivity was obtained for the highest experimented dilution rate (D = 1.0 d−1; i.e., for non N‐limited growth conditions), whereas the highest carbohydrate productivity was obtained at D = 0.67 d−1. We used these results to discuss the nitrogen conditions that optimize NL productivities in the context of biofuel production.

Mesoscale distribution of zooplankton in the Sub-Antarctic Frontal system in the Indian part of the Southern Ocean: a comparison between optical plankton counter and net sampling

Abstract ( Antartic Research (ANTARES) IV Cruise, January–February 1999). Distribution of zooplankton biovolume/biomass in the Sub-Antarctic Frontal system is described for the Indian Ocean sector, northwest of Kerguelen Islands, with a comparison of two types of sampling techniques: WP2 net and optical plankton counter (OPC). During the ANTARES IV Cruise (January–February 1999), three zones were sampled: the Polar Frontal Zone (PFZ), south of the Sub-Antarctic Front; the Frontal Zone between the Sub-Tropical Front; and the Agulhas Front and the Sub-Tropical Zone, north of the main stream of the Agulhas Return Current. Copepods were the dominant group in all net samples regardless of the area considered. Maximum values were recorded in the PFZ with both net catches and OPC. A strong contrast in terms of population structure and biomass was observed between the zone south of the Sub-Antarctic Front and the zones to the north. The patterns of the zooplankton biovolume distribution by size confirmed this relationship. Biovolume 2D maps showed in this south area a more patchy distribution than in the other areas where a vertical gradient dominated. The OPC data are in fair agreement with net sample data and are spatially heterogenous in both size structure and biomass.

Variation of lipid classes among organs of the Northern krill Meganyctiphanes norvegica, with respect to reproduction.

Lipid content and class in the Northern krill Meganyctiphanes norvegica (digestive gland, stomach, gonad, fat body, abdomen) was investigated and correlated with sex and reproductive stage. Ready to spawn females, have high lipid content in ovaries, while in males and spent females, the major site of lipid deposits was the digestive gland, followed by the fat body. These differences among spawning and spent females are indicative of strong interactions between the ovary and digestive gland and the ovary and fat body during vitellogenesis. Triacylglycerols (TAG) were the major neutral lipid class with high levels in the digestive gland. The major phospholipid was phosphatidylcholine (PC) particularly in the muscular tissue of the abdomen. Phosphatidyl-ethanolamine (PE) and -serine-inositol (PS-PI), were present at intermediate levels. Reproductive males were depleted in TAG and diacylglycerols (DAG) in the digestive gland, gonad and fat body, and had 4 times lower cholesterol in the gonad than ready to spawn females. Furthermore, ready to spawn females had in the ovary higher amounts of TAG, DAG and phospholipids (PC, PE, PS-PI) than spent females. Linear relationships between lipid content and main lipid class (TAG, PC, PE, PS-PI) in different fractions of males and ready to spawn females showed that: (1). TAG was stored for both sexes in all cephalothorax fractions with highest values in the digestive gland and ovary fluid; (2). PC was accumulated for both sexes in the fat body and the gonad with a higher slope for females, with the highest values in the ovary fluid and in the abdomen of males and that (3). PS-PI was stored only in the ovary and abdomen of mature females. These results are discussed in terms of the strategy developed by Meganyctiphanes norvegica to allocate lipids to the next generation for optimised embryogenesis.

Lipid composition of the euphausiids Euphausia vallentini and Thysanoessa macrura during summer in the Southern Indian Ocean

Two species of euphausiids (Thysanoessa macrura Sars and Euphausia vallentini Stebbing) from the Southern Indian Ocean were analysed for their lipid content, fatty acid and fatty alcohol composition, in relation to developmental stage (T.m) and sex (E.v). Lipid composition showed wax esters and triacylglycerols as main neutral lipids for T. macrura and E. vallentini respectively. Allometric relationships between lipid class and total lipids indicated that changes in total lipids were size dependent and mainly related to wax esters in T. macrura and polar lipids (both phosphatidylcholine or PC and phosphatidylethanolamine or PE) in E. vallentini. No difference in lipid composition could be shown for male and female E. vallentini, while sampling location, developmental stage and sex were significantly influential in T. macrura. In this latter species wax esters displayed relatively similar fatty acid and alcohol composition in both juvenile and female stages, whereas a striking difference could be seen between females and juveniles in terms of polar lipids with a very low PE content in females. Male and female E. vallentini showed little differences in fatty acid structure with the exception of PC in females, which were low in 22:6n-3. The trophic status of these two species was established using multivariate discriminant analysis, which indicated for E. vallentini a degree of omnivory similar to E. superba, while these T. macrura appeared more omnivorous than individuals collected in other areas of the Southern Ocean. Clustering of polar lipid composition suggested a link between differences in PC fatty acid and the post-spawn stage of the female of E. vallentini collected. The same probably applies for the changes in PE recorded for T. macrura females.

Spatio-temporal distribution of mesozooplankton in a sub-Antarctic coastal basin of the Kerguelen Archipelago (southern Indian Ocean)

Seventeen surveys were performed fortnightly from January to December 1990 and covered a grid of 16 stations in the large Morbihan Bay at Kerguelen Island. Temperature and salinity were measured at four or five standard depths. Chlorophyll, carbon and nitrogen contents of surface water particulate matter were determined along with mesozooplanktonic biomass and abundance of copepods. The dominant species (90%) of mesozooplankton wasDrepanopus pectinatus, and two other taxa,Oithona spp. andCalanus simillimus, accounted for 8.6 and 0.4% respectively. A Multiple Correspondence Analysis was used to determine the influence of abiotic (temperature and salinity) and biotic factors (potential food supply) on the spatio-temporal distribution of copepods and, more specifically, on that of the growth stages ofD. pectinatus. No specific hydrological features were found. The distribution of copepods was fairly homogeneous in the whole bay while the quantitative changes were influenced by the seasonal rhythm. Only a slight trend of increasing values for the biotic or abiotic parameters and the neriticD. pectinatus was observed from the central to the inner back area of the bay.