The role of faecal material in the particulate organic carbon flux in the northern Humboldt Current, Chile (23{degrees}S), before and during the 1997-1998 El Nino

Sedimentation rates of faecal material, phytoplankton and microzooplanktonand production rates of faecal material from crustaceans andpelagic tunicates were estimated during the austral summer andwinter 1997, and summer 1998, in the northern Humboldt Current(23°S, off Antofagasta, Chile). Sampling periods coveredpre-El Niño (January 1997) and El Niño 1997–98(July 1997 and January 1998). Samples were collected using floatingsediment traps deployed at 65, 100, 200 and 300 m depth in oceanicand coastal areas. Sedimentation rates during January 1997 were,on average, 152 ± 23 and 85 ± 57 mg C m^–2day^–1 at 65 and 300 m depth, respectively. During July,these rates averaged 93 ± 56 mg C m^–2 day^–1at 65 m depth and 35 ± 12 mg C m^–2 day^–1at 300 m depth, while in January 1998 they were 98 and 109 ±37 mg C m^–2 day^–1 at 65 and 200 m depth, respectively.Recognizable faecal material made up the bulk of the sedimentingmatter, accounting for 8 ± 5% (n = 14), 31 ± 26%(n = 16) and 8 ± 5% (n = 5) of the average total organiccarbon recorded from all sediment trap samples collected duringJanuary and July 1997 and January 1998, respectively. However,at300 m depth, the contribution of recognizable faecal materialto total sedimented organic carbon increased to 43 ±33% (n = 4) during July 1997. The remaining sedimenting particlesconsisted mainly of tintinnids, crustacean exuviae, heterotrophicdinoflagellates (both thecated and athecated) and diatom cells.During this study, we estimated that only a minor fraction (average± SD = 5 ± 8%) of the copepod faecal materialproduced within the photic zone sedimented down to 300 m depth,suggesting an efficient recycling within the overlaying watercolumn. On the other hand, an important fraction (47 ±30%) of the euphausiid faecal strings was collected in the 300m depth trap, suggesting that this material would enhance thedownward flux of particulate organic matter (POC). POC fluxesto 65 and 300 m depth traps were in the range of 4–20%and 3–8% of the estimated primary production during thewhole study period. It is postulated that the overall verticalflux of particulates and, in particular, faecal pellets wasdetermined by a combination of three factors. The first wasthe composition of the zooplankton assemblages in the studyarea. When the dominant group was calanoid copepods, their faecesseemed to contribute poorly to the vertical flux of particulates.On the other hand, when the dominant group was euphausiids,a significant proportion of their faecal material was collectedin the sediment trap located at 300 m depth. The second wasthe relatively high abundance of cyclopoid copepods from thegenera Oncaea, Corycaeus and Oithona, which are reported tofeed on aggregates of phytodetritus and faecal pellets producedby calanoid copepods, suggesting that they may act as a naturalfilter to sedimenting particulates. The third was the compositionand size spectrum of the phyto- and microzooplankton assemblageswhich are potential food sources for the meso- and macrozooplankton.These factors were partially modulated by both the 1997–1998El Niño and seasonality.     

Spatial distribution of small pelagic fish larvae in the Gulf of California and its relation to the El Nino 1997-1998

Sardinops caeruleus larvae were almost absent during the ElNiño 1997–1998, when the sea surface temperature(SST) increased by 4°C. After the event, these larvae appearedin high abundance between 18 and 22°C SST. Engraulis mordaxlarvae were recorded in high abundance during the El Niño(17 and 25°C SST) and afterwards (14 and 22°C). Duringthe event, they tended to concentrate close to Isla Angel deLa Guarda and Isla Tiburón, the coldest zone. The ElNiño conditions affected the spawning of S.caeruleusmore than E.mordax, with the latter more adapted to strong environmentalchanges.     

Ammonia-oxidizing beta-proteobacteria from the oxygen minimum zone off northern Chile

The composition of ammonia-oxidizing bacteria from the beta-Proteobacteria subclass (betaAOB) was studied in the surface and upper-oxycline oxic waters (2- to 50-m depth, approximately 200 to 44 microM O(2)) and within the oxygen minimum zone (OMZ) suboxic waters (50- to 400-m depth, < or =10 microM O(2)) of the eastern South Pacific off northern Chile. This study was carried out through cloning and sequencing of genes coding for 16S rRNA and the ammonia monooxygenase enzyme active subunit (amoA). Sequences affiliated with Nitrosospira-like cluster 1 dominated the 16S rRNA gene clone libraries constructed from both oxic and suboxic waters. Cluster 1 consists exclusively of yet-uncultivated betaAOB from marine environments. However, a single clone, out of 224 obtained from the OMZ, was found to belong to Nitrosospira lineage cluster 0. To our knowledge, cluster 0 sequences have been derived from betaAOB isolated only from sand, soil, and freshwater environments. Sequences in clone libraries of the amoA gene from the surface and upper oxycline could be grouped in a marine subcluster, also containing no cultured representatives. In contrast, all 74 amoA sequences originating from the OMZ were either closely affiliated with cultured Nitrosospira spp. from clusters 0 and 2 or with other yet-uncultured betaAOB from soil and an aerated-anoxic Orbal process waste treatment plant. Our results reveal the presence of Nitrosospira-like betaAOB in both oxic and suboxic waters associated with the OMZ but with a clear community shift at the functional level (amoA) along the strong oxygen gradient.     

Larval dynamics of the sand crab, Emerita analoga, off the central Oregon coast during a strong El Nino period

Populations of the sand crab, Emerita analoga, are well establishedon the California coast. However, populations in Oregon occursporadically and appear to be restocked by larvae drifting northfrom California. Due to increased northward transport in winterduring El Niño events, we would expect that larval recruitmentto northern populations should be higher during these periods.This hypothesis was tested by comparing larval abundances betweennon-El Niño and El Niño years, 1997 and 1998,respectively. In 1997, larval abundances and seasonality weresimilar to historical data from 1969–71. First zoeal (Z1)stage larvae were present during these summers, but in low numbers,indicating that E.analoga spawned off the coast of Oregon. In1998, strong larval recruitment from the south occurred, demonstratedby a large number of fifth zoeal stage (Z5) larvae collectedoff Oregon in April and May. The large numbers of Z1 found insummer 1998 indicate that the Z5, which arrived from Californiaearlier that spring, successfully recruited to adulthood andreproduced. These data indicate that Oregon E.analoga populationsare probably dependent on larvae travelling from Californiain the Davidson Current. It follows that this species couldbe used as an indicator of coastal current fluctuations suchas those seen during El Niño events.     

Functional diversity of marine macrobenthic communities from sublittoral soft-sediment habitats off northern Chile

Benthic communities show changes in composition and structure across different environmental characteristics and habitats. However, incorporating species biological traits into the analysis can provide a better understanding of system functioning within habitats. We compare the functional diversity of macrobenthic communities from a contrasting shallow (15 m) and deep (50 m) sublittoral soft-sediment habitats in northern Chile, using biological traits analysis. Our aim was to highlight the biological characteristics responsible for differences between habitats and the implications for ecosystem functioning. Trait analysis showed that the deep habitat was restricted in providing functionally important biogenic structure and bioturbation and supports less diverse feeding-related energy pathways. The shallow habitat is characterized by more diverse energy pathways and a higher potential for matter exchange through bioturbation. We provide support to the predictions of transfer of energy from the benthos to upper trophic levels in the shallow, which is characterized mainly by normoxia and little organic matter content in the sediment. In the deep habitat, characterized by hypoxia and more organic matter, energy appears to be transferred to microbial components. We suggest that trait analysis should be added to the traditional approaches based on species diversity, because it provides indicators of ecosystem stress.     

Characterization by PCR of Vibrio parahaemolyticus isolates collected during the 1997-1998 Chilean outbreak

Between November 1997 and April 1998, several human gastroenteritis cases were reported in Antofagasta, a city in the north of Chile. This outbreak was associated with the consumption of shellfish, and the etiologic agent responsible was identified as Vibrio parahaemolyticus. This was the first report of this bacterium causing an epidemic in Chile. V. parahaemolyticus was the only pathogenic bacterium isolated from patient stools and from shellfish samples. These isolates were analyzed by polymerase chain reaction (PCR) amplification of the pR72H gene, a species-specific sequence. Based on the pR72H gene amplification pattern, at least three different isolates of V. parahaemolyticus were found. Two isolates (named amplicons A and C) generated PCR products of approximately 400 bp and 340 bp respectively, while another type of isolate designated B, did not generate a PCR product, regardless of which method of DNA purification was used. Sequence analysis of the amplicons A and C shows that they have an 80 bp and 183 bp conserved region at the 5' end of the gene. However, both isolates have different sequences at their 3' terminus and are also different from the pR72H sequence originally reported. Using this PCR assay we demonstrated that these three isolates were found in clinical samples as well as in shellfish. The warm seawater caused by the climatological phenomena "El Ni?o" perhaps favored the geographic dispersion of the bacterium (bacterial bloom) occurring in Antofagasta that occurred during that time of year.     

Distribution and foraging behaviour of the Peruvian Booby (Sula variegata) off northern Chile

The Peruvian Booby (Sula variegata) is one of the most numerous guano bird species in the Humboldt Current. We used a combination of data logger deployment, at-sea observations and colony-based work to investigate the foraging and diving behaviour, as well as the at-sea distribution and food choice of Peruvian Boobies breeding at Isla Pajaros, northern central Chile. Birds foraged in the vicinity of Isla Pajaros. They performed short foraging trips, varying between a few minutes and several hours. Average foraging range was 17.1 ± 3.5 km and diving took place between 2 and 10 km off the coast. Birds dived to depth of up to 10 m with an average depth of 4.3 ± 1.5 m. By at-sea observations as well as by data logger deployment, we identified important foraging areas close to the coast, especially around protruding points. The birds’ diet was highly variable between years, with Inka scad (Trachurus murphyi) and anchovy (Engraulis ringens) being the overall dominant prey species. Despite limited sample sizes, this paper presents first results about the foraging behaviour of Peruvian Boobies at the largest breeding colony in northern Chile. This information is essential for a better understanding of the effects of environmental changes as well as for the implementation of conservation measures.     

Denitrification, nitrate turnover, and aerobic respiration by benthic foraminiferans in the oxygen minimum zone off Chile

Population density, nitrate turnover, and oxygen respiration of benthic foraminiferans were investigated in the oxygen minimum zone (OMZ) off the Chilean coast. Live foraminiferans were found predominantly in the upper 3 mm of the sediment, and the nitrate accumulating species Nonionella cf. stella and Stainforthia sp. dominated with a combined standing stock of 2.0 × 10⁶ Rose Bengal stained specimens m^− 2. The rate of denitrification in cells of N. cf. stella analyzed with nitrous oxide microsensors during acetylene inhibition was 84 ± 33 pmol C individual^− 1 d^− 1. Multiplied with the standing stock of N. cf. stella and Stainforthia sp. this yielded a minimum benthic denitrification rate of 173 µmol N m^− 2 d^− 1 by foraminiferans. Foraminiferal denitrification, which seemed to account for almost all benthic denitrification at the investigated site will be overlooked by most conventional methods measuring benthic denitrification. Compared to the denitrification rates, the potential rates of nitrate accumulation and oxygen respiration by N. cf. stella were an order of magnitude higher (864 pmol N individual^− 1 d^− 1 and 760 ± 87 pmol C individual^− 1 d^− 1, respectively), which seems an adaptation to the infrequent availability of nitrate and oxygen in the sediment surface.     

Nauplii of the copepod, Calanus pacificus, off southern California in the El Nino winter-spring of 1992, and implications for larval fish

Female and naupliar Calanus were rarer in winter and springof 1992, an El Niño period, than in 1989. The ratio ofabundances, nauplius III (NIII) female^-1 (a composite surrogatefor reproduction and survival), did not differ between yearsand was not consistently correlated with concurrently measuredchlorophyll (a measure of food). Survival from egg to NIII wasnegatively correlated with biomass of macrozooplankton (a measureof predators). Survival from NIV to NVI was less variable thanthat of younger states; it was poorer in winter of 1992 thanin winter of 1989, but did not differ in median magnitude betweenthe two springs, although it was least variable spatially inspring of 1992. Survival of these feeding stages was not correlatedwith chlorophyll. The relative ranty of nauplii was equivalentto a reduction of 30% in this source of food for larval fish.     

Nanoheterotroph grazing on bacteria and cyanobacteria in oxic and suboxic waters in coastal upwelling areas off northern Chile

The vertical distribution and abundance of microbial assemblagesand the grazing of nanoheterotrophs upon prokaryotes in oxicand suboxic waters were examined in two coastal upwelling areasoff northern Chile where a shallow Oxygen Minimum Zone (OMZ)is characteristic. Prokaryotic prey included bacterioplanktonand cyanobacteria (Synechococcus); both displayed a bimodaldistribution, with abundance maxima above and within the upperOMZ. Flagellates numerically dominated the nanoplankton andwere mostly concentrated in the oxic layer. Mean ingestion ratesof cyanobacteria by nanoflagellates (vacuole content method)ranged from 0.2 to 1.1 cells flagellate^–1 h^–1 andmean consumption rates (34–160 cells mL^–1 h^–1)were four times higher in the oxic layer. With the selectiveinhibitors technique, specific grazing rates on bacteria werelow (<0.1 h^–1) and consumption did not control bacterialproduction in the surface layer but did so in the suboxic layer(accounting for >100% of bacterial production). With thesame method, the specific grazing rate on cyanobacteria rangedbetween zero and 0.23 h^–1 with no clear differences betweenoxygen conditions; prey growth and production were always higherthan the grazing pressure (accounting for <17% of cyanobacterialproduction). The impact of grazing by nanoheterotrophs in regulatingthe production of prokaryotes in oxic and suboxic waters inthis region is discussed.     

Epidemiology of human hydatidosis in the IX Region of la Araucania, Chile 1991-1998

In the region of la Araucania, located in the south of Chile (37 degrees 24'-39 degrees 34' South lat.), the economy is based in cattle raising and agriculture. The cases of human hydatidosis, mortality and notification registered in la Araucania 1991-1998 are reviewed. The rate for hydatid disease is 38.5 per 100,000 population. Liver (47.1%) and lung (40.7%) were the most affected viscera. Days in hospital, 11-20 the 56.1%. Disease in female is 51.1%. The mortality rate is 0.8 per 100,000, the most frequently affected county is Imperial. The age group 15-44 years is the most affected. The notification rate is 8.74 per 100,000 population. The rate for hydatid disease is higher than the rate of notification, then necessarily there exist a subnotification. We think that it is necessary to investigate the serological prevalence of hydatidosis in the population of the IX Region Araucania.     

Ontogenetic vertical distribution and diel migration of the copepod Eucalanus inermis in the oxygen minimum zone off northern Chile (20-21{degrees} S)

The vertical and ontogenetic distribution, and diel verticalmigration (DVM), of Eucalanus inermis in relation to the strongvertical gradient in oxygen concentration associated with anintense oxygen minimum zone (OMZ) were studied at a coastalarea off northern Chile (20–21° S). A close relationshipbetween the abundance of the whole copepod population and lowoxygen waters was found, with most developmental stages remainingnear the base of the oxycline (30–80 m) and within theupper zone of the OMZ (30–200 m). All stages performedDVM but not at all the stations, mainly between the 30–60and 60–200 m strata; a small fraction (<20%) appearedin the surface layer (0–30 m) mostly at night. This strategyof movement would result in a better utilization of food resourcessince the strong physical and chemical gradients at the baseof the oxycline and upper OMZ boundary might serve as a siteof particle accumulation. A secondary fluorescence peak was,in fact, found at all the stations, coinciding with minimaldissolved oxygen (DO, <1 mL O₂ L^–1) at the base ofthe oxycline or in the upper OMZ boundary. The relevance ofthe biogeochemical flux involved in this diel migration patternwas assessed by calculating the potential active input of carbonand nitrogen from the upper layers into deeper the OMZ.     

External respiration function of young male residents of northern Europe during the annual cycle

In young men (19.0 ± 0.9 years of age), the following parameters were studied during the annual cycle: the tidal and minute lung volumes, vital and forced vital capacities of the lungs, expiratory and inspiratory reserve volumes, 0.5-and 1-s forced expiratory volumes, and Tiffenau index. Young men working under the conditions of the North (62°N) proved to have deeper breathing; the minute volume and vital capacity of their lungs were increased. Analysis of the lung volume during the annual cycle demonstrated changes in most parameters studied (except the expiratory reserve volume and Tiffenau index). The maximum values of the lung volumes were recorded in the cold time of the year (from November to April), whereas the minimum values were observed in the warm time (from May to September).     

Epidemiology of Echinococcus granulosus in Arbil province, northern Iraq, 1990-1998

During the period 1990-1998, 99 cases of human cystic hydatidosis (12.4 cases per year) were surgically treated at the two main hospitals in Arbil province, northern Iraq, and from this the human occurence for the province was estimated to be 2 per 100,000 inhabitants. In the same area, 1270 sheep, 550 goats and 320 cattle were examined at slaughter for hydatid cysts and prevalence rates were found to be 15.0%, 6.2% and 10.9%, respectively. A decreasing tendency in livestock prevalences was found towards the end of the study period. As in humans, most of the hydatid cysts in livestock were located in the liver. Fertility of sheep cysts, i.e. those containing protoscoleces, was found to be significantly higher (64%) than that of goats (35.7%) and cattle (29.8%). The percentage of fertile cysts containing viable protoscoleces varied between 63 and 82% in the livers and between 72 and 79% in the lungs of the different animal species. A total of 97 stray dogs were examined post-mortem in the years 1991, 1992 and 1998, and Echinococcus granulosus worms were found in the intestines of 48 dogs (49.5%). High worm burdens (> 1000) were observed in 37% of the dogs, medium worm burdens (200-1000) in 41%, and low worm burdens (< 200) in 22%. In 1998, the prevalence of canine echinococcosis (24.3%) was found to be significantly lower than in 1991 (70.4%) and 1992 (60.6%). The prevalence of human hydatidosis did not differ significantly over the years, but the study confirmed that hydatidosis is endemic in northern Iraq, and that housewives, labourers and farmers appear to be at the greatest risk of infection.     

Human remains from Blombos Cave, South Africa: (1997-1998 excavations)

The Middle Stone Age (MSA) layers at Blombos Cave contain abundant bifacial Still Bay points, formal and ad hoc bone artefacts, and an intentionally incised bone piece. These artefacts add weight to arguments that some aspects of modern human behavior developed earlier in sub-Saharan Africa than elsewhere. Four human teeth were recovered from the MSA strata at Blombos during the 1997-1998 excavations. Two are heavily worn deciduous teeth, and two are incomplete permanent premolar crowns. The Blombos di(1)is comparatively large in relation to modern African homologues, falling within the lower part of the observed Neandertal range. The dm(1)and P(3)are comparable to modern teeth and smaller than most Neandertal crowns. The premolars preserve horizontal circum-cervical striae that suggest palliative toothpick use. The di(1)evinces labial scratches that resemble neither the "cutmarks" that have been observed on Neandertal incisors, nor the striae that have been recorded on modern human teeth.     

Egg production rates of eight calanoid copepod species during summer 1997 off Newport, Oregon, USA

Measurements of hydrography, water transparency, chlorophyll (Chl) a and egg production rates (EPRs) by females of Calanus marshallae Frost, Calanus pacificus Brodsky, Eucalanus californicus Johnson, Epilabidocera longipedata Sato, Pseudocalanus mimus Frost, Centropages abdominalis Sato, Acartia longiremis Lilljerborg and Paracalanus parvus (Claus) were estimated at weekly intervals between 17 July and 2 September 1997. Production of eggs was determined in 24 h incubations to examine the effects of environmental variability on EPR, to detect the possibility of food limitation of EPR, and to evaluate the hypothesis that growth rates of females are size dependent. During the study, an anomalous downwelling event occurred, possibly in response to the 1997 El Niño, which allowed us to determine how El Niño events affect EPRs of coastal copepods. The larger copepods Calanus marshallae, Calanus pacificus and Centropages abdominalis showed the highest egg production and specific growth rates during the period of active upwelling (18 July-13 August, water temperatures 8-13C, Chl a concentration 4.7-16.2 l^-1 and water transparency 3-5 m). After 27 August, the 1997-98 El Niño arrived off Oregon, creating a downwelling situation. Upwelling winds ceased, the thermocline intensified, temperature and transparency increased (to >18°C and 16 m), and Chl a declined to <2 g l^-1. Densities of the common coastal species declined greatly as well. Paracalanus parvus became the dominant species, and Eucalanus californicus, Epilabidocera longipedata and Corycaeus anglicus became common in our samples. EPRs for the larger boreal copepods (Calanus and Centropages) declined greatly during El Niño;; the smaller copepods, Pseudocalanus mimus, A.longiremis and Paracalanus parvus, showed low but relatively constant egg production and specific growth rates during both upwelling and downwelling events. Over the entire study period, only three species produced eggs at its maximum rate (24-28 eggs female^-1 day^-1), Pseudocalanus mimus which averaged 4 eggs day^-1. All other species had EPRs that were two or five times below their maximum EPR. Thus, EPRs were not related to body size, contrary to our expectations. Hatching success was variable among species. Values as low as 20-40% were found for all species at least once during the study period, suggesting that occasionally a substantial portion of egg production may not be viable.      

Susceptibility to selected coagulase-negative chemotherapeutic agents of methicillin resistant staphylococci isolated from clinical materials in 1997-1998

The study was aimed at assessment of the sensitivity of methicillin-resistant coagulase-negative staphylococci isolated from clinical material in 1997/1998 to selected chemotherapeutic agents. The investigated material comprised 96 methicillin-resistant coagulase-negative staphylococci from hospital and ambulatory infections isolated during the period from April 1997 to May 1998. Species affiliation was determined by classical identification methods and commercial diagnostic tests for identification of staphylococci. Methicillin resistance was determined by agar disk-diffusion method and screening. Sensitivity to chemotherapeutics was determined by agar disk-diffusion method and agar dilution methods. All the investigated strains were sensitive to nitrofurantoin, furazolidone and vancomycin. To teicoplanin--the second glycopeptide antibiotic--84% strains were sensitive, whereas the percentages of resistant and moderately sensitive strains amounted to 5.2% and 10.4%, respectively. 85% and 82% of coagulase-negative staphylococcal strains were sensitive to fusidic acid and mupirocin. Considerable differences were noted with respect to sensitivity to aminoglycoside group antibiotics. About 35% of strains were sensitive to gentamicin, and 90% sensitive to netilmicin. Ca. 40% of coagulase-negative staphylococci were resistant both to cotrimoxazole and trimethoprim, which, in view of 98% resistance to the second component of cotrimoxazole, may be associated with the activity of only one of the components of the drug--trimethoprim.     

Prevalence of scrapie infection in Great Britain: interpreting the results of the 1997-1998 abattoir survey

An accurate estimate of the prevalence of scrapie infection in the Great Britain (GB) sheep flock is essential when assessing any potential risk to human health through exposure to sheep transmissible spongiform encephalopathies (TSEs). One method for assessing the prevalence is to sample sheep intended for human consumption using a diagnostic test capable of detecting infected animals prior to the onset of clinical signs. An abattoir survey conducted in Great Britain in 1997-1998 tested brain samples from 2809 apparently healthy sheep of which none was found to be positive for scrapie by histopathology or immunohistochemistry (IHC) although 10 were positive for scrapie-associated fibrils (SAF). Subsequently, the tonsils from a subset of the animals sampled were examined using IHC, one of which tested positive. To interpret these results we use a likelihood-based approach, which accounts for the variation in the prevalence of infection with age and test sensitivity and specificity with stage of infection. Combining the results for all of the diagnostic tests yields an estimate of the prevalence of scrapie infection in the GB sheep flock of 0.22% (95% confidence interval: 0.01-0.97%). Moreover, our analysis suggests that all of the diagnostic tests used are very specific (greater than 99%). Indeed, only SAF detection yields a specificity estimate of less than 100%, which helps to account for the high number of samples found to be positive for SAF.     

Grazing impact of copepod assemblages and gravitational flux in coastal and oceanic waters off central Chile during two contrasting seasons

In evaluating carbon flux in coastal and oceanic waters offcentral Chile (~36°S), the grazing pressure by copepod size-assemblagesand the gravitational flux from the surface layer were estimatedduring two contrasting seasons: spring upwelling (October 1998)and winter (July 1999) periods. Grazing pressure upon phytoplanktonbiomass was small (<5%) during both periods at all stations.It was, however, an important proportion of primary productionat the coast during the spring (17–43%) but minimal inwinter (<5%) while, at the oceanic station, it was significantduring both seasons (13–46%). Similarly, the downwardflux of particulate organic carbon was a significant percentageof primary production at the coast during the spring (31%) andless so in winter (15%); at the oceanic station, it was a higherfraction during both seasons (46–47%). Copepod pelletswere only a minor component of the faecal flux, suggesting thatthey were rapidly recycled in the water column. The seasonaldifferences in these carbon fluxes in the coastal upwellingzone were related directly to the larger quantities of organicmatter in the water column during the spring upwelling period,including higher chlorophyll a concentrations and primary productionrates. During the winter, the particulate organic carbon appearsto be mainly recycled in the water column. In the oceanic zone,in contrast to what is expected, a large percentage of the primaryproduction appears to be exported during both periods, thoughconsumption by copepod assemblages was also important, suggestingsignificant horizontal carbon export from the coast to the openocean.