Biological Oceanography · Biological Oceanography SAYED Z. EL-SAYED Department of Oceanography...

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Biological Oceanography SAYED Z. EL-SAYED Department of Oceanography Texas A&M University When Eltanin made her maiden voyage to the icy waters of the Antarctic during Cruise 4 in July 1962, the main objective of the marine biological program was to increase the knowledge of the biol- ogy and ecology of the seas surrounding Antarctica. In this respect, the ship's mission did not differ much from those of her celebrated predecessors Belgica, Pourquoi Pas?, Meteor, Gaus, and Dis- covery. Any reviewer of Eltanin's accomplishments in biological oceanography between 1962 and 1972 certainly must be impressed by the rapid evolution from exploratory studies to more sophisticated and highly integrated programs designed to study the total antarctic ecosystem. In fact, there were two distinct phases. The first began with Cruise 4 in the Drake Passage and ended with Cruise 36, more than halfway around the antarctic continent in the Tas- man Sea, in November 1968. This period is charac- terized by (a) accumulation of comprehensive and representative collections of marine fauna and flora, (b) study of distribution and biogeography of the antarctic and subantarctic flora and fauna with em- phasis on relationships of the biota to nonantarctic regions, (c) use of physical and chemical oceano- graphic data in the study of the ecology of the marine organisms, (d) initiation of studies on the physiological mechanisms of adaptation in antarctic invertebrates (mainly crustacea) to persisting low temperatures, afl(1 (e) systematic study of the bio- logical productivity of the waters surrounding Ant- arctica. During that phase, the biological investigations were but one aspect of Eltanin's numerous and broad scientific programs, which included marine geology and geophysics, physical oceanography, marine chemistry, upper atmospherics, and meteor- ology. These multidisciplinary programs typified the early cruises. The conflicting interests of the biological and physical programs, together with the urgency of studying the structure and function of the ecosys- tem, ushered in Eltan in's secon(l phase of biological investigation. The phase began with Cruise 38 (March to May 1969) . Together with two other biological cruises (46 and 51) , this phase was a landmark in the history of biological exploration of the seas surrounding Antarctica. The latest cruise in this trilogy ended in February 1972 with the com- pletion of an integrated study of the biology, chem- istry, and physical oceanography of the Ross Sea. Unlike the earlier phase, with its emphasis on sur- veying, reconnaisance, and intensive collecting, the 1969-1972 period featured interrelated system- oriented studies of the antarctic ecosystem as a functioning unit. This phase has witnessed (a) documentation and extension of data for the stand- ing crop and primary production of phytoplankton on a seasonal basis in the area from lO°W. to 100E. and from 140°S. into the pack-ice, (b) assessment of the standing stocks and biomass of the secondary producers (zooplankton) together with seasonal variability, (c) description of the distribution of organic materials throughout the water column, in- chiding all dissolved and particulate materials, (d) understanding of microbial activity in the minerali- zation processes in antarctic waters, (e) determina- tion of the biochemical composition of planktonic and nektonic organisms with respect to transfer of complex lipids in the antarctic food chain, (f) use of carbon-13 and -14 ratios to trace the pathways of carbon as it passes through the various trophic levels, (g) acquisition of experimental data on stress measurements of the response of phytoplank- ton (in situ and in vitro) to changing light and temperature, and (h) collection of data on trophic efficiency estimates and respiration, energy require- ments, and energy content of zooplankton to pro- vide baseline data on energy flow. The following pages summarize the research in these two phases. On account of the multitude and diversity of the investigations, the accomplishments made in each of the main components of the marine ecosystem are discussed separately. The earlier phase l'hytoplankton During Cruises 13, 14 and 15 (May to December 1964) the late Paul Burkholder began a study of the distribution of the standing crop of phyto- plankton (using chlorophyll a method) and primary production of the South Pacific Ocean. Wood (1967) studied the vertical distribution of the phytoplankton in the Bransfield Strait and at a series of stations between South America and New Zealand. He found that the maximum phytoplank- May-June 1973 93

Transcript of Biological Oceanography · Biological Oceanography SAYED Z. EL-SAYED Department of Oceanography...

Page 1: Biological Oceanography · Biological Oceanography SAYED Z. EL-SAYED Department of Oceanography Texas A&M University When Eltanin made her maiden voyage to the icy waters of the Antarctic

Biological OceanographySAYED Z. EL-SAYED

Department of OceanographyTexas A&M University

When Eltanin made her maiden voyage to the icywaters of the Antarctic during Cruise 4 in July1962, the main objective of the marine biologicalprogram was to increase the knowledge of the biol-ogy and ecology of the seas surrounding Antarctica.In this respect, the ship's mission did not differmuch from those of her celebrated predecessorsBelgica, Pourquoi Pas?, Meteor, Gaus, and Dis-covery.

Any reviewer of Eltanin's accomplishments inbiological oceanography between 1962 and 1972certainly must be impressed by the rapid evolutionfrom exploratory studies to more sophisticated andhighly integrated programs designed to study thetotal antarctic ecosystem. In fact, there were twodistinct phases. The first began with Cruise 4 in theDrake Passage and ended with Cruise 36, more thanhalfway around the antarctic continent in the Tas-man Sea, in November 1968. This period is charac-terized by (a) accumulation of comprehensive andrepresentative collections of marine fauna and flora,(b) study of distribution and biogeography of theantarctic and subantarctic flora and fauna with em-phasis on relationships of the biota to nonantarcticregions, (c) use of physical and chemical oceano-graphic data in the study of the ecology of themarine organisms, (d) initiation of studies on thephysiological mechanisms of adaptation in antarcticinvertebrates (mainly crustacea) to persisting lowtemperatures, afl(1 (e) systematic study of the bio-logical productivity of the waters surrounding Ant-arctica.

During that phase, the biological investigationswere but one aspect of Eltanin's numerous andbroad scientific programs, which included marinegeology and geophysics, physical oceanography,marine chemistry, upper atmospherics, and meteor-ology. These multidisciplinary programs typifiedthe early cruises.

The conflicting interests of the biological andphysical programs, together with the urgency ofstudying the structure and function of the ecosys-tem, ushered in Eltan in's secon(l phase of biologicalinvestigation. The phase began with Cruise 38(March to May 1969) . Together with two otherbiological cruises (46 and 51) , this phase was alandmark in the history of biological explorationof the seas surrounding Antarctica. The latest cruisein this trilogy ended in February 1972 with the com-

pletion of an integrated study of the biology, chem-istry, and physical oceanography of the Ross Sea.Unlike the earlier phase, with its emphasis on sur-veying, reconnaisance, and intensive collecting, the1969-1972 period featured interrelated system-oriented studies of the antarctic ecosystem as afunctioning unit. This phase has witnessed (a)documentation and extension of data for the stand-ing crop and primary production of phytoplanktonon a seasonal basis in the area from lO°W. to 100E.and from 140°S. into the pack-ice, (b) assessmentof the standing stocks and biomass of the secondaryproducers (zooplankton) together with seasonalvariability, (c) description of the distribution oforganic materials throughout the water column, in-chiding all dissolved and particulate materials, (d)understanding of microbial activity in the minerali-zation processes in antarctic waters, (e) determina-tion of the biochemical composition of planktonicand nektonic organisms with respect to transfer ofcomplex lipids in the antarctic food chain, (f) useof carbon-13 and -14 ratios to trace the pathways ofcarbon as it passes through the various trophiclevels, (g) acquisition of experimental data onstress measurements of the response of phytoplank-ton (in situ and in vitro) to changing light andtemperature, and (h) collection of data on trophicefficiency estimates and respiration, energy require-ments, and energy content of zooplankton to pro-vide baseline data on energy flow.

The following pages summarize the research inthese two phases. On account of the multitude anddiversity of the investigations, the accomplishmentsmade in each of the main components of the marineecosystem are discussed separately.

The earlier phase

l'hytoplankton

During Cruises 13, 14 and 15 (May to December1964) the late Paul Burkholder began a study ofthe distribution of the standing crop of phyto-plankton (using chlorophyll a method) andprimary production of the South Pacific Ocean.Wood (1967) studied the vertical distribution ofthe phytoplankton in the Bransfield Strait and ata series of stations between South America and NewZealand. He found that the maximum phytoplank-

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ton distribution occurred at a depth of 100 metersin the BransfIeld Strait and between the surface and30 meters in the Pacific sector.

Beginning with Cruise 18, the Texas A&M groupembarked on one of the most extensive investiga-tions of primary productivity ever undertaken inthe Southern Ocean (fig. 1) . This investigation in-cluded the systematic study of spatial, seasonal, andyear-to-year variations of the standing crop of phy-toplankton, species composition and species divers-ity of the phytoplankters, nutrient chemistry (in-cluding phosphates, silicates, nitrates, nitrites, par-

ticulate and dissolved organic carbon) , and a studyOf the hydrographic conditions affecting the produc-tivity of these waters. The results of this investiga-tion (Balech et al., 1968) underscored the conspic-uous regional differences in the productivityrameters of the Southern Ocean and pointed to thestriking differences between the productivity of theoceanic and neritic regions (El-Sayed, 1970) . Thedata substantiated the hypothesis that the richnessof the antarctic waters is real only with regard to thecoastal and inshore regions, and not with regard tothe oceanic regions. As a result of these productivity

Figure 1.Location of stations occupied in circumantarctic waters where primary productivity studies were carried out aboard the Eltanin, USCG

Glacier, ARA 'General San Martin, and Capitan Canepa (modified from El-Sayed, 1970).

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measurements, the early and persistent views re-garding the proverbial fertility of the antarcticwaters have given way to more realistic estimates ofthe productivity of that region of the world ocean.Walsh (1969) compared the phytoplankton stand-ing cropin the Southern Ocean with that of theStrait of Florida. His data suggest that despite ap-parent high production during the austral summer,the overall production of the Antarctic and temper-ate and subtropical areas may be at least of thesame order of magnitude.

Zooplankton

Extensive studies of the zooplankton populationsbegan with the Lamont antarctic plankton programduring Cruise 8. The objective was to study thedistribution and abundance of the zooplanktonpopulation between the surface and 2,000 meters.Special collecting devices included (a) it multipleplankton sampler that collects at depths of 500 to250 meters, 250 to 100 meters, and 100 meters tothe surface (luring an oblique tow, and (b) a bathy-pelagic plankton sampler calibrated to sample from1,000 to 500 meters. Since Cruise 15, a second bathy-pelagic plankton sampler has been collecting atdepths from 2,000 to 1,000 meters. Zooplankton wascollected systematically and fairly continuously(luring the early cruises. Portions of the samplescollected were sent to the Smithsonian Oceano-graphic Sorting Center for sorting and distributionof the organisms among the specialists throughoutthe world (Fehlmann, 1971; Smithsonian Oceano-graphic Sorting Center, 1969).

Typical of the zooplankton studies conductedduring this phase are those of Chen and Ericson(1967) who studied the vertical distribution of themajor groups of holoplanktonic gastropods andcorrelated them with different water masses atdifferent depths. Hillman (1969) studied theplankton samples from the Pacific sector of theAntarctic and the Scotia Sea and determined theseasonal quantitative distribution of the ostracodsfrom 0 to 1,000 meters. Caldwell (1966) investi-gated the distribution of pelagic tunicates based onthe material collected in the Drake Passage and ad-jacent areas, and Be (1969) mapped the distribu-tion of the planktonic foraminifera in the antarcticand subantarctic waters.

An important aspect of the zooplankton investi-gations (luring the early cruises was the estimationof the zooplankton standing crop. From the collec-tions made in 1963 to 1965 aboard Eltanin between750 and 185°W. (an area sparsely studied by Dis-covery) , Hopkins (1971) assayed the standing cropof zooplankton using 202-micron closing nets in

the upper 1,000 meters. He was able to show thatthe principal groups were copepods, chaetognaths,and euphausiids, with the copepods alone constitut-ing 67.3, 68.8 and 70.1 percent of the biomass inthe antarctic, subantarctic, and convergence zonewaters, respectively. Total biomass per square meterin the upper 1,000 meters averaged 2.67, 2.58 and2.96 grams (Try weight in the antarctic, the subant-arctic, and the convergence zones, respectively, withlongitudinal or seasonal variability being difficult todetect. Hopkins also noted a relatively large fractionof carnivores, which generally exceeded 20 percentof the total zooplankton standing crop. His dataimply that on the basis of Slobodkin's (1960) find-ings, standing crop cannot always be used reliablyto mirror net trophic efficiency.

Fishes

Except for DeWitt (1970) , the study of the ich-thyofauna was limited (luring the early cruises.Bussing (1965) studied the collections of midwaterfishes taken along the western coast of SouthAmerica in the Peru-Chile Trench. He found that83 percent of the species apparently do not crossthe boundary centered at 20°S. that separates Pa-cific equatorial water masses from subantarcticwater masses. The other 17 percent were found wellon both sides of the boundary. In observing thecharacter of the midwater fish fauna of the RossSea, DeWitt (1970) found that the striking decreasein the temperature of deep water in the Ross Seais the likely explanation for the absence there ofpelagic fishes of the southern ocean. DeWitt in-dicated that there is essentially no midwater faunain the Ross Sea and that Plenragramma antarcticumis the only species occurring in great numbers in theRoss Sea. DeWitt (1971) makes an excellent con-tribution to our knowledge of antarctic coastal anddeep water benthic fishes; a good portion of thesedata were collected aboard Eltanin.

Benthos

Marine biologists from the University of SouthernCalifornia and other institutions in the UnitedStates have filled huge gaps in our knowledge ofthe distribution and ecology of benthic fauna(Hartman 1964, 1966). Representative of the ben-thic fauna studies (luring the early cruises are Foster(1967), who summarizes Harvard University'sbrachiopods (about 10,000 specimens) taken fromantarctic waters and off Antipodes and MacquarieIsland, Herb's (1971) study of the distribution ofrecent benthic foraminifera in the Drake Passage,Kott's (1969, 1971) investigations of the ascidians,

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and Clark's (1970) study of the collection of as-teroids made by Eltanin at depths from 167 to 4,686meters off the New Zealand coast.

The fauna of the trenches bordering South Amer-ica were studied by Menzies (1963). Since he wasnot able to show that antarctic deep-sea specieswere found in the trenches, he tentatively concludedthat the fauna of these trenches are cosmopolitanand mostly from low latitudes. Menzies and George(1969) studied the patterns of distribution of thedeep-sea antarctic isopod crustacea; they showed atendency of the shallow-water genera to penetrate(lee1) in the abyss and a simultaneous tendency ofthe abyssal genera to emerge into shallow water.These two phenomena—polar submergence andpolar emergence—remain clearly distinguishablefrom the standpoint of taxonomic perception of thegenera involved. George and Menzies (1968) pro-vided evidence to show a seasonal breeding cycleof the abyssal isopods, with the peak breedingperiod limited to 4 months of the year, July toNovember. One of the many Eltanin firsts was thediscovery by Rosewater (1970) in the AtlanticOcean off southeastern South America of a smallspecimen belonging to the subgenera Neopilina col-lected from i depth of 1647 to 2044 meters. Thisdiscovery may indicate a widespread distribution ofthis abyssal group in the World Ocean.

Other research

Other research during the early cruises includesEltan in's ornithological research program, whichwas begun by the Dominion Museum (Wellington,New Zealand) in January 1965 to study the seasonaldistribution and relative abundance of seabirds inthe Antarctic and subantarctic. About 45 species ofseabirds were obtained. A rookery of over 10 mil-lion chinstrap penguins was found on ZavodovskiIsland (Harper, 1966)

Airborne insects in the Antarctic were collectedby screening air while the ship was at sea. Of the679 specimens collected, 454 were Diptera and 86were Homoptera (Holzapfel et al., 1970)

It is fitting that the earlier phase culminated inBe et al. (1969), where all the marine invertebrates- planktonic foraminifera, porifera, nemertea,brachirpoda, bryozoa, sipuncula, benthic mollusca,pycnogonida, planktonic ostracoda, amphipoda,crinoidea, holothuroidea, echinoidea, asteroida, op-hiuroidea, and ascdiaca - are discusse(l and theirdistributions mapped.

The later cruises

Cruise 38, which centered on study of metabolic

processes in the southern ocean, blazed the trailfor the two subsequent integrated biological cruises.These cruises were needed because, although wehave a fairly reasonable knowledge of the composi-tion, abundance, and distribution of the maincomponents of the antarctic ecosystem, we still lackinformation about the relationship between thetrophic levels and the flow of energy through thatecosystem. The time had come when studies of onecomponent or another, which typified the Eltanininvestigations between 1962 and 1967, could notreliably provide coherent and correlated data forcomputer simulation and prediction. What wasneeded, El-Sayed (1971) pointed out, was a team ofinvestigators whose correlated efforts would havethese objectives: (a) determination of the trophiclevels and estimation of the biomass tied up in eachlevel, (b) determination of pathways and flow ofnutrient materials and energy and estimation offlow rates, and (c) measurement of the physicaland chemical milieu of the community.

The accomplishments of Eltanin Cruise 38, whoseactivities centered around the study of the metab-olic processes of the living organisms in antarcticand subantarctic waters, are discussed in McWhin-nie (1973).

Cruise 46 (November 1970-January 1971) cul-minated a 3-year effort to mount balanced, inte-grated programs in biological and physical ocean-ography together with biochemical and radiationStudies. These programs were designed to solvesome of the outstanding problems of the antarcticmarine ecosystem. Cruise 51 (January 1972-Febru-ary 1972) was the second integrated biology cruisewith the objective of contributing to our knowledgeof the functional relationships and the energy inthe trophic hierarchy of the antarctic marine eco-system (fig. 2) . The contributions made by thesethree cruises are summarized below.

Solar radiation available to the euphotic zone

Since photosynthesis by different species of phyto-plankton is dependent on light at different parts ofthe spectrum, a knowledge of the spectral distribu-tion of solar radiation received at the surface—whichIS a function if the altitude of the sun, the turbidityof the atmosphere, and the amount afl(1 type ofclouds—is basic to ;I understanding of pri-mary productivity. During Cruises 16 and 51, effortswere directed toward determining the amount andquality of light made available in the water column(Franceschini, 1971, 1972). The approach employ-ed two similar sets (one set measuring the down-welling stream: the other, the upwelling flux) offour hemispheric sensors to measure continuously

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PHYTOPLANKTONPRIMARY PRODUCTIONCHLOROPPIYLL, , B PHAEO-PFWTtIBIOMASS & SPECIES COMPOSITIONNET PLANKTON B NANNOPLANKTONLIP[)SCALORIC VALUESCHEMICAL COMPOSITION. AT PZOOSPO4RC B SAPROPHYTIC FUNGIMARINE BACTERIA

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the radiation fluxes in four wavebands a short dis-tance above the surface. Franceschini (1972) wasable to show that the quality of the energy madeavailable for photosynthesis differs from that ofthe incident light. For a detailed account of studieson solar radiation, see Franceschini (1973).

Ecological studies of phytoplankton

Marine phytoplankton investigations were di-rected toward increasing our understanding of theecology of the primary producers and toward abetter knowledge of the dynamics of the lowertrophic levels of the food chain. Such investigationsare essential to the study of the functional relation-ships and the energy flow through the trophiclevels of the ecosystem. In studying these trophicrelationships, it is imperative to know the amountof carbon fixed by the phytoplankton. This was

accomplished by measuring primary organic pro-duction using the in situ carbon-14 uptake tech-nique instead of the simulated in situ methods per-formed during the earlier cruises (El-Sayed, 1969).In the later cruises, expanded study was made ofthe standing crop of phytoplankton in the upper300 meters of the water column and of the contri-bution of nannoplankton to both the standing cropand primary productivity. During Cruise 51, thenannoplankton were found to contribute about 75percent of the phytoplankton standing crop and 80percent of the primary production. (Fay, unpub-lished manuscript)

Marine phycomycetes

Although studies of marine fungi began in theearlier cruises (Fell, 1968) , it was not until thelater cruises that concerted efforts were made to

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Figure 2. Aspects of the structure and function of antarctic ecosystems as studied during Cruise 51.

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study their occurrence and distribution. Bahnwegand Sparrow (1972) studied the occurrence anddistribution of the marine fungi from south of NewZealand and Australia to the antarctic continent.Primary emphasis was given marine phycomycetes,although the occurrence of yeasts and filamentoushigher fungi (molds) also was recorded. Their pre-liminary results indicate that "molds" are infre-quent in oceanic waters, a finding that correlateswell with the results obtained by the above authorsduring Cruise 46 in the southern Indian Ocean.However, in contrast to the results of Cruise 46,yeasts also were rare. All of the phycomycetes foundsouth of the polar front are somewhat similar toI)crmocystidium sp. sensu Goldstein and Moriber(1966) in that they have neither rhizoids nor mo-bile zoospores.

Carbon pathways

The pathways of carbon were traced through theantarctic ecosystem using natural variations in car-bon and hydrogen isotopes (Sackett and Brooks,1972). With knowledge of carbon-13/carbon-12and deuterium/hydrogen ratios in inorganic corn-pounds and in marine organisms, it is possible topredict the pathways of carbon through the marineecosystem beginning with carbon dioxide in theatmosphere and ending with the final members ofthe food chains.

Heterotrophic potential of marine microorganisms

The rate of heterotrophic activity was measuredby kinetic analysis of the assimilation and respira-tion of carbon-14 labeled glucose, acetate, and var-ious amino acids. Morita and his associates fromOregon State University are interested in measur-ing the extent of microbial participation in themineralization process in antarctic waters. The rateat which this process proceeds has important im-plications in determining the route of energy flowthrough the biosphere as well as determining thepotential inorganic nutrient recycling rates associ -ated with microbial activity (Morita et al., 1972)

Particulate and dissolved organic matter; planktonchemistry

During Cruises 46 and 51, detailed vertical pro-files of dissolved and particulate carbon, nitrogen,and phosphorus were taken at 35 stations betweenAustralia, New Zealand, and Antarctica. Thesedeterminations, in conjunction with inorganic ni-trate, phosphate, and silicate concentrations have

shown that these biologically important nutrientsare not strikingly different in deep water from otheroceans (Carlucci et al., 1972) . Thus, water sinkingat the polar front does not contribute a high con-tent of dissolved organic matter to intermediatewaters of the Pacific Ocean. The investigation isimportant in evaluating in situ oxygen concentra-tions and rates of microbial oxidation with time indeep water. The Scripps Institute of Oceanographygroup also studied the distribution of particulateorganic carbon (oc), nitrogen (P0N), and phos-phorus (o) at the 35 stations mentioned above.The total particulate organic carbon values in theeuphotic zones ranged from 30 to 200 microgramsof carbon per liter. In all profiles the particulateorganic carbon decreased rapidly at depths between200 and 500 meters, below which it ranged about2 to 10 micrograms of carbon per liter. The carbon/nitrogen ratios of the particulate matter rangedfrom about 4 to 10, without any significant variationwith depth. Most of the C/N ratios are close to 5.0,in sharp contrast to water of other oceans where theC/N is close to 10.0. The poc/pop ratio varies withdepth from about 20 to 80; this ratio is significantlylower than those found in lower latitudes, wherethe ratio often is between 50 and 200.

Vitamins B 11 thiamine, and biotin

Although a number of reports cite the distribu-tion of dissolved vitamin B 1 , thiamine, and biotinin the world ocean, there was no adequate informa-tion on vitamin distribution in antarctic watersbefore Cruise 46. Water samples were taken atvarious depths during Cruise 46 and Cruise 51. Re-sults of the earlier cruise indicate that thiaminewas probably not important in the productivity ofthe waters since it was found in only a few samples,including those from deeper water. Biotin appearedto be produced byby the phytoplankton, since it wasdirectly correlated with high phytoplankton popu-lations in the upper waters from most stations. Thedirect involvement of B 12 in the ecology of phyto-plankton is more obvious in waters of Antarcticathan in most areas of the world oceans.

Also during Cruise 51 the Scripps investigatorscollected samples for determination of mercury andcadmium, carbon-l4 and tritium content, andchlorinated hydrocarbons. These samples includewater, ice, surface films, sediments, marine aerosols,and a number of organisms representing varioustrophic levels in the antarctic food chain (Holm-Hansen, personal communication) . This work isessential in assessing variations with time of thesechemical components in the Southern Ocean, withrespect to rates of transfer into antarctic waters

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from their primary source in the northern hemis-phere, as well as their flux rate through the antarc-tic food chains.

Physiology of antarctic zooplankton

Respironietric study of pelagic invertebrates wascarried out (luring Cruise 51 (McWhinnie and Kir-chenberg, 1972). Species studied included the eu-phausiids Euphausia superba, E. crystellorophias,and E. triacantha; the amphipods Parathemistogaudichaudii and P/iron ima sp., the copepod Gala-noides acutus, the chaetognath Eu/trohnia sp. andthe tunicates Salpa sp. and Pyrosoma sp. Oxygenconsumption values for most animals were generallyequivalent to those obtained during Cruise 46 inthe southern Indian Ocean (McWhinnie and Ur-banski, 1971). McWhinnie and her students alsostudied the use of soluble nutrients by various zoo-plankton. They had reportedreported previously that ant-arctic amphipods, studied in the Pacific sector dur-ing Cruises 17 to 19, utilize soluble nutrients (Mc-Whinnie and Johanneck, 1966) as do a number ofspecies studied in the Indian Ocean sector duringCruise 46 (McWhinnie and Urbanski, 1971) . Thedata obtained (luring Cruise 51 support earlier in-terpretations concerning a direct route of absorp-tion of soluble organic compounds by zooplankton(McWhinnie and Kirchenberg, 1972) . For a detail-

ed account regarding the physiology and metabolismof the marine organisms in the Antarctic, see Mc-Whinnie (1973).

crystailorophias. The differences between these twoeuphausiids were found to be due to the presence ofwaxes in E. crystallorophias but not in E. superba(Bottino, 1972; also personal communication).

Clearly, synthesis of the above-mentioned studiescarried out during the later cruises, namely pro-grams of solar radiation and spectral quality oflight, biological activity and chemical compositionof plankton, primary productivity and phytoplank-ton standing crop, metabolism and energy turnoverof zooplankton, role of marine fungi, pathways ofcarbon through the food chains, lipid metabolism,heterotrophic bacteria, should provide us with oneof the most comprehensive pictures so far of thefunctioning of the antarctic marine ecosystem.

There is more work to do. Some of the major ant-arctic marine resources (e.g., krill) are threatenedby commercial exploitation, and marine biologistsmust develop guidelines, based on hard data, forthe wise use of these resources within the frameworkof the ecosystem concept. The tragedies of over-exploitation of the plaice fishery in the North Sea,the California sardine, and, lest we have forgotten,the antarctic baleen whales are glaring examples ofpoor management of natural resources. Whilemarine biologists lament the layup of the ship,there is no denying that it was Eltanin that main-tained U. S. leadership in antarctic marine biologyin the 1960s. During her short service in the causeof science, Eltanin has earned her rightful placeamong the celebrated polar research vessels.

ReferencesLipid metabolism

Studies since 1966 have clarified various aspectsof the lipid composition and metabolism of antarc-tic organisms. Early investigations by Jeffrey et al.(1966) showed that in many of the organisms phos-pholipids are the most abundant lipid class. Fromthese investigations it also was apparent that marinephospholipids are much richer than marine tria-cylglycerols in long-chain highly unsaturated fattyacids (HUFA) . Consequently, the HUFA of marinefish and mammals seem to originate mainly fromthe phospholipids in organisms of the lower trophiclevels rather than from the triaclyglycerols of thesame organisms (Bottino, personal communica-tion) . The positional distribution of HUFA in thelipids of zooplankion, euphausiids, and fish wasfound to be similar in all three groups. Studies dur-ing Cruise 51 showed that whereas the fatty acidsprofile of a particulate species, E. superba, for ex-ample, is quite constant, it can differ markedly fromthe fatty acid profile of another species such as E.

Bahnweg, G., and F. K. Sparrow, Jr. 1972. Marine phycomy.cetes; occurrence south of New Zealand and the Ross Sea.Antarctic Journal of the U.S., VII (5) : 177-178.

Balech, E., S. Z. El-Sayed, G. Hasle, M. Neushul, and J . S.Zaneveld. 1968. Primary productivity and benthic marinealgae of the Antarctic and Subantarctic. Antarctic MapFolio Series, 10. 12 p. 15 plates.

Be, A. W. H., H. Boschma and T. P. Lowe, J . S. Bullivant,E. \V. Dawson, J . H. Dearborn and J . A. Rommel, R. K.Dell, S. J . Edmonds, H. B. Fell and S. Dawsey, H. B. Fell,T. Holzinger, and H. Sherraden, M. W. Foster, S. R. Geigerand C. Brahm, J . W. Hedgpeth, N. S. Hillman, D. E.Hurley, V. M. Koltun, P. Kott, D. L. Pawson, A. Ross and\V. A. Newman, D. F. Squires. 1969. Distribution of se-lected groups of marine invertebrates in waters south of35S. latitude. Antarctic Map Folio Series, 11. 44 p. 29plates.

Bottino, Nestor R. 1972. Fatty acid exchange among trophiclevels of the Ross Sea: phytoplankton, copepods, and eu-phausiids. Antarctic Journal of the U.S., VII (5) : 178-179.

Russing, W. A. 1965. Studies of the midwater fishes of thePeru-Chile trench. Antarctic Research Series, 5: 185-227.

Caldwell, M. C. 1966. The distribution of pelagic tunicates,family salpidae, in antarctic and subantarctic waters. South-ern California Academy of Sciences Bulletin, 65 (1) : 1-16.

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Carlucci, A. F., 0. Holm-Hansen, and P. M. Williams. 1972.Distribution and composition of particulate and dissolvedorganic matter in antarctic waters. Antarctic Journal of theU.S., VII(5): 181.

Chen, C., and D. B. Ericson. 1967. Holoplanktonic Gastropodain the southern oceans. Antarctic Journal of the U.S.,11(5): 260.

Clark, H. E. S. 1970. Sea-stars (Echinodermata: Asteroidea)from Eltanin Cruise 26, with a review of the New Zealandasteroid fauna. Victoria University, Wellington. ZoologicalPublication, 52. 33 p.

DeWitt, H. H. 1970. The character of the midwater fish faunaof the Ross Sea, Antarctica. In: Antarctic Ecology, (M. W.Hoidgate, ed.) , vol. 1. New York, Academic Press. p. 305-314.

DeWitt, H. H. 1971. Coastal and deep-water benthic fishes ofthe Antarctic. Antarctic Map Folio Series, 15. 10 p., 5 plates.

El-Sayed, S. Z. 1969. Ecological studies of antarctic marinephytoplankton. Antarctic Journal of the U.S., IV(5): 193-194.

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Fay, R. R. Significance of the nannoplankton in the produc-tivity of the Ross Sea, Antarctica. (unpublished manu-script)

Fehlmaun, H. Adair. 1971. USARP activities at the Smith-sonian Oceanographic Sorting Center. Antarctic Journal ofthe U.S., VI (6) : 250-251.

Fell, J . W. 1968. Distribution of antarctic marine fungi. Ant-arctic Journal of the U.S., III (5); 157.

Foster, M. W. 1967. A summary of Harvard University'sbrachiopod studies on Eltanin Cruise 27. Antarctic Journalof the U.S., 11(5): 192.

Franceschini, G. 1971. Observations of net solar radiation foroceanic biological study. Antarctic Journal of the U.S.,VI (5) : 157-158.

Francescluini, G. 1972. Spectral components of solar radiationavailable to the euphotic zone. Antarctic Journal of theU.S., VII (5) : 175-176.

Franceschini, G. 1973. Solar radiation: USNS Eltanin, 1962-1972. Antarctic Journal of the United States, VIII (3) : 108.

George, R. Y., and R. J . Menzies. 1968. Species of storthyn-gina (isopoda) from the Antarctic with descriptions of sixnew species. Crustaceana, 14(3): 275-301.

Goldstein, S., and L. Moriber. 1966. Biology of a problematicmarine fungus, Dermocystidium sp. I. Development andcytology. Archiv fur Mikrobiologie, 53: 1-11.

Harper, P. C. 1966. A New Zealand ornithologist on Eltanin.Antarctic (Wellington) , 4 (8) : 389-390.

Hartman, 0. 1964. Polychaeta errantia of Antarctica. Antarc-tic Research Series, 3. 131 p.

Hartman, 0. 1966. Polychaeta myzostomidae and scdentariaof Antarctica. Antarctic Research Series, 7. 158 p.

Herb, R. 1971. Distribution of recent bethonic foraminiferaiii the Drake Passage. Antarctic Research Series, 17: 251-300.

Hillman, N. S. 1969. Ontogenic studies of antarctic pelagicostracoda. Antarctic journal of the U.S., IV (5) : 189-190.

Holzapfel, E. P., D. M. Tsuda, and J . C. Harrell. 1970. Trap-ping of airborne insects in the antarctic area (Part 3).Pacific Insects, 12 (1) : 133-156.

Hopkins, T. L. 1971. Zooplankton standing crop in the Pa-cific sector of the Antarctic. Antarctic Research Series, 17:347-362.

Jeffrey, L. M., N. R. Bottino, and R. Reiser. 1966. The dis-tribution of fatty acids classes in lipids of antarctic Eu-phausiids. Antarctic Journal of the U.S., 1(5): 209.

Jitts, H. R., and 1). J . Carpenter. Eltanin Cruise 38. III. Re-sponses of phytoplankton photosynthesis to variations inlight and temperatures. (unpublished manuscript)

Kott, P. 1969. Antarctic Ascidiacea. Antarctic Research Series,13. 239 p.

K()tt, P. 1971. Antarctic Ascidiacea II. Antarctic ResearchSeries, 17: 11-82.

Menzies, R. J . 1963. General results of biological investigationson the deep-sea fauna made on the USNS Eltanin (u5ARP)

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Menzies, R. J . , and R. Y. George. 1969. Polar faunal trendsexhibited by antarctic isopod crustacea. Antarctic Journalof the U.S., IV(5): 190-191.

\IcWhinnie, M. A., and R. Johanneck. 1966. Utilization ofinorganic and organic carbon compounds by antarctic zoo-plankton. Antarctic Journal of the U.S., I (5) : 210.

McWhinnie, M. A., and R. J . Kirchenberg. 1972. Physiologyof antarctic zooplankton with special reference to crusta-ceans. Antarctic journal of the U.S., VII (5) : 176-177.

McWhinnie, M. A., and R. J . Urbanski. 1971. Absorption ofsoluble organic compounds by polar marine zooplankton.Antarctic Journal of the U.S., I(S): 210.

McWhinnie, M. A. 1973. Physiology and biochemistry: USNSEltanin, 1962-1972. Antarctic Journal of the U.S., VIII (3)101.

Morita, R. Y., R. P. Griffiths, and S. S. Hayasaka. 1972. He-terotiopinc potential of antarctic marine microorganisms.Antarctic Journal of the U.S., VII (5) : 180-181.

Rosewater, J . 1970. Monoplacophora in the South AtlanticOcean. Science, 167 (3924) : 1485-1486.

Sackett, W. M., and J . M. Brooks. 1972. Stable carbon iso-tope variations in the antarctic marine ecosystem. AntarcticJournal of the U.S., VII (5) : 179-180.

Slohodkin, L. B. 1960. Ecological energy relationships at thepopulation level. American Naturalist, 194 (876) : 213-236.

Smithsonian Oceanographic Sorting Center. 1969. The Smith-sonian Oceanographic Sorting Center. The Science Teacher,March 1969: 29-31.

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Walsh, J . J. 1969. Vertical distribution of antarctic phyto-plankton II. A comparison of phytoplankton standing cropsin the southern ocean with that of the Florida Strait.Limnology & Oceanography, 14 (1) : 86-94.

Wood, E. J . F. 1967. Antarctic phytoplankton distribution.Antarctic journal of the U.S., 11(5): 190.

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