On the place of origin of deep-sea lsopods

Biogeographic and phylogenetic data on the deep-sea isopod family Ilyarachnidae (Paraselloidea) document its origin and evolutionary radiation in the deep sea. The distribution of eyes among paraselloidean families suggests an in situ evolution for all those families which have primarily deep-sea distributions. Present-day distributions of paraselloidean isopods provide no hints to their ultimate sources in shallow water. These findings contrast to recent suggestions in the literature that the deep-sea isopod fauna has been derived from shallow Antarctic waters.     

Origin of Antarctic Isopoda (Crustacea,Malacostraca)

An analysis of the horizontal and the vertical zonation of the Antarctic Isopoda, combined with knowledge of the geological history of Antarctica and isopod phylogeny, revealed that the isopod family Serolidae and subfamily Arcturinae are likely to have evolved from ancestors that inhabited a cold-temperate Gondwanian province. Antarctic species of other families, such as the Munnopsidae, Nannoniscidae, Desmosomatidae and Ischnomesidae, are likely to have evolved from deep-sea ancestors. It is deduced that emigration of South Patagonian species into the Southern Ocean, although possible, probably did not occur very often. Evolutionary phenomena such as continental-drift vicariance, radiation of species on the continental shelf of Antarctica, and active migration, including submergence and emergence mechanisms are discussed.     

Foraminiferan (Protozoa) epizoites on Arctic isopods (Crustacea) as indicators of isopod behaviour?

A total of 38 219 specimens representing 63 species of marine isopods (Crustacea) from deep and shallow Arctic waters were studied in a search for epizoic foraminifers (Protozoa). Foraminifers occurred on 21 species, and their frequency was generally low. A total of 290 foraminifer individuals were found, of which 289 belonged to Cibicides wuellerstorfi, C. refulgens and Cibicides spp. (juveniles) (Cibicidae), while only a single individual belonged to Cornuspira sp. (Cornuspiridae). The foraminifers were most frequent on species of the families Munnidae, Ischnomesidae (suborder Asellota) and on Gnathia stygia (suborder Gnathiidea), but were totally absent from the asellote families Janiridae, Haploniscidae, Nannoniscidae and from the suborder Epicaridea. The foraminifers were mainly located on the legs (Munna acanthifera), the anterior part of the body (Haplomesus quadrispinosus, heteromesus frigidus), or on the head (G. stygia adults). The epizoic foraminifers occur mainly on epibenthic isopods, which do not or only rarely clean themselves. The foraminifers are known to prefer elevated substrata, and in this the habitat of the isopods and the foraminifers coincide. The size of individual isopods was not related to the presence or absence of foraminifers.     

Diet of four species of deep-sea isopods (Crustacea: Malacostraca: Peracarida) in the South Atlantic and the Southern Ocean

The food of four species of asellote isopods (Crustacea, Malacostraca), Haploniscus rostratus, Haploniscus unicornis, Acanthocope galatheae and Betamorpha fusiformis, was evaluated by analysis of their gut contents. The isopods were sampled at several stations on the abyssal plains of Guinea Basin, Angola Basin and Cape Basin (southeast Atlantic), the Weddell Sea abyssal plain and the Antarctic continental slope during the DIVA and ANDEEP expeditions in 2000, 2001 and 2005. While all species had mineral particles in their guts and mucus material was the most frequent food item, the remaining gut contents differed among species. Betamorpha fusiformis fed mostly on phytodetritus, especially in the Southern Ocean basins and ingested along with it whole calcareous foraminifers. Acanthocope galatheae showed some differences in gut contents between basins, but in the Guinea Basin, the contents were to a large extent stercomata, i.e., waste pellets of soft-walled foraminifers, i.e., the Komokiaceae. Indications were that the haploniscids were feeding on detritus and agglutinating foraminifers (stercomata). This indicates spatial differences in food availability for this diverse group of deep-sea isopods and the importance of poorly known foraminiferal groups, like the Komokiaceae, as a food source in the deep sea.     

Temperature and pressure tolerance of larvae of Crepidula fornicata suggest thermal limitation of bathymetric range

The extant deep-sea fauna is thought to result from recolonisation of this environment by shallow-water organisms following climate-driven mass extinctions. Planktonic larval tolerance to high pressure is considered an important preadaptation for successful deep-sea invasion. In this study, the pressure and temperature tolerance of a species without any known confamilial deep-sea relative were assessed for the first time. Early- and late-veliger larvae of the shallow-water species Crepidula fornicata were subjected to a temperature/hydrostatic pressure regime from 5 to 25 °C and from 0.1 to 40 MPa. Although early and late veliger survived pressures equivalent to 2,000 m water depth or greater at all temperatures, decreased larval activity indicated significant sublethal temperature and pressure effects. Reduced larval activity of early veliger at low temperatures suggests that the bathymetric range of this species may be thermally constrained. A mechanistic model is proposed to explain the emerging pattern of ontogenetic shifts in pressure tolerance of shallow-water benthic invertebrates.     

A comparative survey of neuston: geographical and temporal distribution patterns

Neuston samples were collected from 149 stations during a voyage west from Fiji to the Bay of Biscay from August 1979 through November 1980. The neustonic fauna was counted and assigned to 50 species groups, chosen to represent the most numerous animals in the hauls. Analyses of neustonic concentration and population structure showed regional and temporal differences in the fauna. Permanent inhabitants of the surface waters, the euneuston, were most numerous in the open tropical waters of the Central Indian Ocean. In contrast, the temporary neuston, which was composed of juveniles, vertical migrants and the uppermost individuals from deeper populations, attained its greatest concentration in the upwelling region of the Somali Current and at inshore stations around the Indonesian archipelago and in the Gulf of Aden and Red Sea. Heavy rains adversely affected the neuston, possibly by diluting the surface waters. Both neustonic groups were sparsely represented in the cooler waters of the Mediterranean Sea and the eastern Atlantic Ocean. At night considerable immigration occurred into the neustonic zone, and the mean faunal density at dusk was 21 times greater than was found at mid-day. Surface waters were inhabited by the developmental stages of many species, and eggs and juveniles of both vertebrates and invertebrates were consistently present in the hauls.     

Depth-related adaptations, speciation processes and evolution of color in the genus Phyllidiopsis (Mollusca: Nudibranchia)

The nudibranch genus Phyllidiopsis (Phyllidiidae) contains 30 currently recognized species, all of them distributed throughout the tropical Indo-Pacific, eastern Pacific, Northwest Atlantic and Caribbean Sea. Half of the known species of Phyllidiopsis inhabit deep waters, and most of the deep-sea species of the Phyllidiidae belong to this genus. There is no definitive explanation for the high diversity of Phyllidiopsis in the deep-sea or for whether diversity could be related to particular adaptations of this group or to historical events. In light of phylogenetic analysis, several cases of vicariance have been detected in this genus. Apparently two major vicariant events occurred between the tropical Indo-Pacific region and the Atlantic-eastern Pacific area first and subsequently between the eastern Pacific and the Atlantic. Vicariant events could also be involved in producing vertical distributional patterns in a few species of Phyllidiopsis. The scarcity of phyllidiids in the Atlantic Ocean may be explained by historical events, including isolation and subsequent extinction in shallow waters. There is a mimicry species complex in Phyllidiopsis, including several members of a clade that probably acquired this coloration through common ancestry, and also including another unrelated species that probably acquired this coloration through convergent or parallel evolution. There is also a group of white species, lacking any other contrasting colors, that inhabits deep waters. This coloration could constitute an adaptation to the deep-sea environment and not a mimicry complex. In this case, all species acquired this coloration through common ancestry.     

Seasonal variation in the diversity and abundance of pelagic larvae of Antarctic marine invertebrates

Most marine benthic macroinvertebrate species reproduce via a larval phase but attempts to explain the occurrence of different larval strategies (feeding or non-feeding, pelagic or benthic) in different habitats have been largely inconclusive. There have been very few year-round surveys of meroplankton at any latitude and in consequence fundamental data on the diversity, abundance, and timings of larval life history phases are lacking. There has been considerable debate regarding the viability of pelagic larvae in cold waters with highly seasonal primary production but there has been only one year-round study of meroplankton in the Southern Ocean, and that was outside of the Antarctic Circle. We present data from the first year-round survey of meroplankton assemblages at a location within the Antarctic Circle. We surveyed abundances of meroplanktonic larvae over 1.5 year at Rothera Point, West Antarctic Peninsula (67°34′S, 68°07′W). Larvae were collected in monthly diver-towed net samples close to the seabed at 20 and 6 m total water depths at each of three locations and were identified and counted live immediately after sampling. A total of 99 operationally defined taxonomic types representing 11 phyla were recorded but this is likely to be an underestimate of true diversity because of inherent difficulties of identification. Larvae were present in all months of the year and although planktotrophic larvae were more abundant in summer, both feeding and non-feeding types were present in all months. Comparisons of seasonal larval abundances with data from a settlement study at the same sites and from the literature show that larvae of mobile adults settle in summer regardless of developmental type, whereas sessile taxa settle in all seasons. We suggest that this is a consequence of differences in the food requirements of mobile and sessile fauna and that the availability of food for post-larval juveniles is more critical for survival than factors affecting the larval stage itself.     

Effect of hydrostatic pressure and temperature on the activity and synthesis of chitinases of Antarctic Ocean bacteria

The formation of chitinases by psychrophilic and psychrotrophic marine Antarctic bacteria and the activity of these extracellular enzymes were investigated under simulated deep-sea conditions. The formation of the chitinases was affected by hydrostatic pressure of 400 bars. However, the extent of pressure inhibition differed with the bacterial strains tested and was considerably less with the extreme psychrophilic bacteria isolated from sediments of greater depth. Growth of these psychrophilic strains had a moderately barophilic character at 400 bars, whereas growth of the psychrotrophic strains was clearly restricted under simulated deep-sea conditions. With regard to the activity of the extracellular chitinases of various bacterial strains, a relatively uniform response was found. All chitinases were highly barotolerant at near neutral pH and were active up to 1000 bars. Low temperatures reduced their activity but not their barotolerance. A low pH of 5.1 diminished the barotolerance of some chitinases. The results suggest that the indigenous deep-sea bacteria are capable of decomposing chitin settled to or produced in the depth of the Antarctic Ocean.     

Temperature adaptation in strains of the amphi-equatorial green alga Urospora penicilliformis (Acrosiphoniales): biogeographical implications

The temperature requirements for growth and the upper survival temperatures (UST's) of the amiphi-equatorial green alga Urospora penicilliformis collected from several localities within its distribution area between 1986 and 1991 were determined. Ecotypic variation, both with regard to growth ranges and optima and to survival temperatures, was demonstrated. In the polar strains of U. penicilliformis, temperature growth ranges were narrower and the growth optima and UST's were at lower temperatures compared to cold-temperate strains. In particular, the polar strains grew between 0 and 15°C with optimal growth at 0 or 5°C, whereas the cold-temperate isolates grew between 0 and (15) 20°C with almost equal growth rates or a growth optimum between 5 and 15°C. The Arctic strains survived 23 to 24°C, and the Antarctic isolate only 19°C, while the UST's of the cold-temperate isolates were between (24) 25 to 26°C. The data strongly indicate that a cold water history of ca. 3 million yr in the Arctic can be sufficient for changes in the temperature growth ranges and optima as well as for small changes of UST as shown in the Arctic populations of U. penicilliformis. For stronger reduction of upper survival temperatures, longer time periods are necessary as exemplified in the isolate from Antarctica, where low temperatures have existed for at least 14 million yr. The significantly lower UST of the Antarctic strain, points to an early contact of the alga with the cold water of the Antarctic region and may indicate an origin of U. penicilliformis in the Southern Hemisphere. The UST's of the cold-temperate isolates (24 to 26°C) would have allowed a migration across the equator during Pleistocene lowerings of the seawater temperatures in the tropics. Growth, however, would not have been possible during the passage across the equator due to the narrow temperature-growth window. The nature of the geographical boundaries and the control of seasonal development of U. penicilliformis by the temperature conditions in the various geographical regions are discussed in relation to the present local temperature regime.     

Pelagic biogeography of the South Atlantic Ocean

The biogeography of the South Atlantic was investigated using presence/absence data for euphausiids. Records were taken from recent and historic, as well as published and unpublished data sets. The resulting biogeography is the most complete to date and can be usefully compared with the biogeochemical provinces for the region. A total of six biogeographic provinces were identified from similarity analyses of the 246 five-degree grid squares. These correspond to Antarctic, sub-Antarctic, cold temperate, warm temperate (subtropical) and tropical waters, as well as the Agulhas Current. Congruence with the biogeochemistry of the region is good in the south and emphasises the important determinate role of temperature. However, the biogeography fails to identify coastal and tropical biogeochemical provinces. This can be attributed to the fact that while adjoining areas may share many species in common, their assemblages differ in their quantitative composition. This serves to emphasise differences in provincial functioning. Received: 28 May 1997 / Accepted: 15 July 1997     

Phytoplankton-zooplankton relationships in the Western Pacific Ocean and adjacent seas

Regressions of biomass and daily food requirements of herbivorous zooplankton on daily primary production were calculated, using assumptions based on data collected in various sea areas of the western Pacific Ocean and adjacent seas. A regression coefficient (1.470) of calculated herbivorous biomass on observed daily primary production is significantly higher than unity (P<0.01). This indicates that the herbivorous biomass sustained by unit amount of primary production is large in the more productive high latitudes, and small in the less productive tropical sea areas. This is attributed to relatively larger food requirements per unit biomass of the tropical herbivores as compared with those found in cold waters. Despite distinct areal differences in the herbivorous biomass-primary production ratios, the calculated daily food requirement of herbivores was in direct proportion to the daily primary production, when equilibrium had been established between phytoplankton and zooplankton. Under conditions of limited food supplies, the small body size of the tropical herbivores may be advantageous both in reducing the total energy consumption per individual, and in inducing rapid growth and reproduction. Therefore, the low ratio of biomass to primary production in the tropics could beregarded as a result of possible regulation of tropical herbivores to scarce food conditions rather than as evidence of failure of adaptation to such conditions.     

Ecological implications of fecal pellets produced by the Antarctic krill Euphausia superba in the Antarctic Ocean

Organic analyses and electron microscopic observations on fecal pellets produced by the Antarctic krill Euphausia superba Dana showed that krill fed on choanoflagellates, the abundant heterotrophic flagellate in the Antarctic Ocean. Two new pathways of organic materials in the Antarctic ecosystem are proposed: (1) a new food chain including non-living particulate and dissolved organics, and bacteria-choanoflagellate-krill-vertebrate, which coexists with the traditional diatom-krill-vertebrate food chain; (2) non-phytoplanktonic organic materials in surface waters are transferred into choanoflagellates and are transported to deep water as fecal pellets which are still useful as nutrition for other organisms there.     

Life hung by a thread: endurance of Antarctic fauna in glacial periods

Today, Antarctica exhibits some of the harshest environmental conditions for life on Earth. During the last glacial period, Antarctic terrestrial and marine life was challenged by even more extreme environmental conditions. During the present interglacial period, polar life in the Southern Ocean is sustained mainly by large-scale primary production. We argue that during the last glacial period, faunal populations in the Antarctic were limited to very few areas of local marine productivity (polynyas), because complete, multiannual sea-ice and ice shelf coverage shut down most of the Southern Ocean productivity within today's seasonal sea-ice zone. Both marine sediments containing significant numbers of planktonic and benthic foraminifera and fossil bird stomach oil deposits in the adjacent Antarctic hinterland provide indirect evidence for the existence of polynyas during the last glacial period. We advocate that the existence of productive oases in the form of polynyas during glacial periods was essential for the survival of marine and most higher-trophic terrestrial fauna. Reduced to such refuges, much of today's life in the high Antarctic realm might have hung by a thread during the last glacial period, because limited resources available to the food web restricted the abundance and productivity of both Antarctic terrestrial and marine life.     

On the near-surface plankton of the eastern South Pacific Ocean

A study has been made of the plankton of the near-surface water layer (0 to 30 cm) of the eastern South Pacific Ocean in the region lying to the east of the meridian 90°W, between latitudes 5°N and 35°S. This region is influenced by the Peru Current: the current brings water from high latitudes, which results in a decrease in the number of species of the local fauna of copepods of the family Pontellidae, typical of tropical near-surface plankton. Some of the widely tropical and one bicentral species are absent or rare. Least affected by the Peru Current are the waters of equatorial structure in the northern part of the region. Here, 7 species of pontellids were recorded: the widely tropical Labidocera detruncata, Pontella tenuiremis, Pontellopsis regalis, the distant-neritic Pontella danae and Labidocera acuta, the bicentral Labidocera acutifrons, and the neritic Pontellopsis lubbockii. The dominant species among these are L. detruncata and L. acuta. To the west of the convergence, in the southern part of the region, live the southern central species Pontella valida and P. whiteleggei, with Pontellopsis regalis occurring occasionally. In these regions the copepod fauna is frequently dominated by pontellids. To the south of the boundary of the waters of equatorial structure, between the coast of South America and the line of convergence, lies a region most subjected to the effect of waters from high latitudes and of upwellings. It is inhabited by 2 pontellids only: Pontellopsis regalis and Labidocera acutifrons, but they too disappear close to the coast. In this particular region the copepods Calanus australis and Centropages brachiatus are common; they are found in a thicker water layer (0 to 200 m), and are often more abundant than the pontellids.     

Stable isotopes identify age- and sex-specific dietary partitioning and foraging habitat segregation in southern giant petrels breeding in Antarctica and southern Patagonia

We examined the isotopic signatures (δ¹³C, δ¹⁵N) of adult body feathers from southern giant petrels Macronectes giganteus collected at two breeding colonies in Antarctica (Potter Peninsula and Cape Geddes) and one in southern Patagonia (Observatorio Island), as well as in whole blood collected from adults of both sexes at each Antarctic colonies and from chicks at Potter Peninsula. As body feather moult is a continuous process in giant petrels, feathers provide an integrated annual signal of an adult’s diets and foraging habitats. In contrast, the stable isotope values of adult and chick blood are reflective of their diets during the breeding season. We found that sex-specific dietary segregation in adults breeding in Antarctica was notable during the breeding season (blood samples) but absent when examined across the entire year (feather samples). In addition, blood stable isotope values differed between chicks and adults, indicating that adults provision their offspring with a relatively higher amount of penguin and seal prey that what they consume themselves. This finding confirms previous work that suggests that chicks are preferentially fed with prey of presumably higher nutritional value such as carrion. Finally, based on isotopic differences between major oceanographic zones in the Southern Ocean, our data indicate population-specific differences in foraging distribution, with Antarctic populations move seasonally between Antarctic and subantarctic zones, while Patagonian populations likely forage in subtropical waters and in continental shelf habitats year-round.     

Light acclimation states of phytoplankton in the Southern Ocean, determined using photosynthetic pigment distribution

In high-latitude waters such as the Southern Ocean, the primary production of phytoplankton supports the ecosystem. To understand the photo-acclimation strategy of such phytoplankton within cold environments, the vertical distribution profile of photosynthetic pigments was analyzed in the Southern Ocean. Samples were taken along 110°E during the austral summer, and along 150°E and around the edge of the seasonal sea ice of the Antarctic Continent during the austral autumn. Pigment extraction methods were optimized for these samples. The standing crop of chlorophyll a was larger in the region along the edge of the seasonal sea ice than at sampling stations in open ocean areas. Chlorophyll concentration seemed to be dependent on the formation of thermo- and haloclines along the edge of the seasonal sea ice, but not in the open ocean where such clines are less pronounced. The marker pigments fucoxanthin and/or 19′-hexanoyloxyfucoxanthin were dominant at most sampling stations throughout the water column, while other marker pigments such as alloxanthin were quite low. This indicated that diatoms and/or haptophytes were the major phytoplankton in this area. Comparison of the relative ratio of fucoxanthin with that of 19′-hexanoyloxyfucoxanthin allowed some stations to be characterized as either diatom-dominant or haptophyte-dominant. The relative ratio of xanthophyll-cycle pigments (diadinoxanthin plus diatoxanthin) to chlorophyll a was high in surface waters and decreased gradually with depth. This suggests that near the ice edge during summer in the Southern Ocean, both diatoms and haptophytes acclimate to their light environments to protect their photosystems under high-light conditions.     

Magellanic Bryozoa: Some ecological and zoogeographical aspects

This study summarizes observations made on more than 140 species obtained in Southern Chile and on many others collected from the Juan Fernández Islands to the Antarctic Peninsula from 1962 to 1980. The total number of the Magellanic Bryozoa including the author's records and those of the literature reaches 195. These species are: 15 Ctenostomata (7.69%), 46 Cyclostomata (23.59%) and 134 Cheilostomata (68.72%). Seven species are described as new: Arachnopusia admiranda sp. nov., Celleporella chiloensis sp. nov., Electra pilosissima sp. nov., Hippoporina chilota sp. nov., Parasmittina pluriavicularis sp. nov., Smittina ectoproctolitica sp. nov. and Smittina molarifera sp. nov. Typical characteristics of the Magellanic bryozoan fauna are: the Cyclostomata amount to 50% of all the subantarctic and antarctic species, with two endemic families Calvetiidae and Pseudidmoneidae; the Cheilostomata show a low endemism at the generic level compared to the antarctic group; the hippothoan fauna is very diversified with four species having tatiform ancestrulae. The zoarial diversity and the polymorphism of the Magellanic Bryozoa are qualitatively and quantitatively different from that of the Antarctic. Thus the encrusting species make up the main bulk of this bryozoan fauna with at least 110 spp. From a bryozoogeographical point of view the Magellanic Region comprises the following parts that could be considered as provinces: (a) Magellanic: including the southern tip of South America south to Lat. 40°S, plus the Patagonian Shelf and the Falkland Islands. (b) Tristan da Cunha and Gough Island. (c) Kerguelen and the nearby archipelagos in the Indian Sector of the Austral Ocean.     

Breeding and larval distribution of the pteropod Clione limacina in the North Atlantic,Subarctic and North Pacific Oceans

The pteropod Clione limacina (Phipps, 1774) is an arcticboreal, circumpolar species, which is widely distributed in the North Atlantic and Subarctic Oceans; it also occurs in the North Pacific Ocean (in the Oyashio and neighbouring waters) and along the Atlantic coast of North America in the waters of the cold Labrador current to the Cape Hatteras region (35° N). The distribution of C. limacina larvae in the plankton of the Norwegian, Barents and White Seas, the Bear Island-Spitsbergen region of the Greenland Sea, the Newfoundland Grand Bank and the Flemish-Cap Bank region of the North-western Atlantic Ocean, and the Kurile-Kamchatka region of the North-western Pacific Ocean has been studied, and information from literature concerning the reproduction and larval occurrence of the species is summarized. Throughout its distributional are, spawning of C. limacina is characterized by the same general ecological pattern. This species breeds and spawns in all types of water masses occurring within the vertical range which it commonly inhabits — from surface layers to 500 m water depth. In all local populations of the species, the most intensive spawning is correlated with the spring/summer period of annual heating of the local waters, and the highest abundance parallels maximum growth of phytoplankton which serves as food for veligers and early polytrochous larvae. After the end of this period, spawning intensity in all local C. limacina populations declines sharply, but spawning continues at low intensity during the autumn/winter season, being practically continuous throughout the year. Distribution patterns of C. limacina larvae are determined by those of their parental forms (the parental forms spawn in the zones permanently inhabited). The earliest larval stages of C. limacina (veligers) are present predominantly in the upper 100 or 200 m water layer, i.e. in the zone of high phytoplankton abundance. Polytrochous larvae, after becoming predaceous feeders, are distributed throughout the whole water column from the surface to 500 m depth, similar to adult C. limacina. As with the adults, larvae are present (within the species' distribution area) in all types of water masses. Since the beginning of the twentieth century, in the course of the warming of the Arctic Ocean, the southern race of C. limacina (formerly a summer/autumn seasonal invader in the Norwegian Sea) has become a permanent component of the plankton fauna of the Norwegian and Barents Seas in regions influenced by the Norwegian-Northcape Current System.     

Distribution of glycosylglycerols and oligosaccharides in the marine environment and their ecological significance in the deep sea

Determination of low molecular weight carbohydrates in marine environments indicated that 1-O--D-galactosylglycerol, 6-O--D-galactosyl-1-O--D-galacto-sylglycerol, sucrose, laminaribiose and laminaritriose are widely distributed in seawaters, suspended and sinking particles, and sediments in coastal as well as in deep-sea waters [e.g. Mikawa Bay and Sagami Bay, Kumano Nada (offshore Japan), and northwest North Pacific Ocean, Bering Sea and Antarctic Ocean: collections during 1978–1984]. Identification of these glycosylglycerols and oligosaccharides in algal cells such as a flagellate (Olisthodisus luteus), blue-green algae (e.g.Trichodesmium sp.) and a diatom (Reptocylindrus denicus) strongly suggests that these sugars are photosynthetically produced by algae in the euphotic zone and are then rapidly transported to the deep sea as sinking particles which can be collected by sediment-trap experiments. The rapid decay rate of low molecular weight carbohydrates by microorganisms suggests that the transported sugars provide energy substrates for microorganisms living in the deep sea.Please address all requests for reprints to Dr. Handa at Nagoya University