OCEANOLOGY, English Translation, VOL. 35, NO.

5, APRIL 1996
Russian Edition: SEPTEMBER { OCTOBER 1995

Characterization of epipelagic ecosystems of the Pacic

Ocean on the basis of the satellite and eld
observations: The plankton stock
in the epipelagial
E. A. Shushkina, M. E. Vinogradov, S. V. Sheberstov, N. P. Nezlin,
and V. I. Gagarin
P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow

Abstract. The Pacic Ocean was divided into zones according to surface chloro-
phyll concentration determined from data of the Nimbus 7 satellite. The areas,
occupied by the waters of seven distinct chlorophyll concentration levels, were
computed. Experimental evidence for the upper 200-m layer, derived from about
300 stations of the P. P. Shirshov Institute of Oceanology, Russian Academy of
Sciences ships, was used to compute geometric mean biomass and to make stock
estimates of phytoplankton, bacteria, protozoa, and mesoplankton for each season
and the annual average for each of the chlorophyll concentration levels. For the
whole Pacic Ocean, this resulted in an annual average stock of 134  106 tons
for phytoplankton; 86  106 tons for bacteria; 26  106 tons for protozoa; and
18:5  106 tons for mesoplankton. The seasonal stock variations were shown to be
insignicant for all of the listed groups. The reason is that in winter the stocks
in the temperate zones decrease, while the areas of oligotrophic waters reduce.
The opposite pattern occurs in spring and summer. The seasonal stability of the
plankton stock, and, consequently, the metabolic process in uence by far the special
features of the carbon cycle in the ocean.

Introduction tem parameters. Quantitative features of these rela-

In earlier studies [Vinogradov et al., 1992, 1993, 1995]
we examined the feasibility of evaluation of dierent
ecosystem parameters and their variability in the upper
200-m layer of the ocean using satellite determinations
of chlorophyll concentration in the surface layer z from
CZCS color scanner data measured at wavelengths of
443, 550, and 670 nm at the Nimbus 7 satellite in 1978{
1986, and experimental evidence collected in the course
tions have been obtained by Vinogradov et al. [1992,
1993, 1995].
Combining satellite and ground truth experimental
evidence makes possible to determine more reliably the
mass of organic matter in diverse groups of plankton or-
ganisms in various regions and in the ocean as a whole.
It becomes possible to trace changes in their stocks and
total content of organic matter in the upper 200-m pro-
ductive layer of the ocean both in biological (climatic
of eld studies on research vessels of the P. P. Shirshov
Institute of Oceanology, Russian Academy of Sciences
(IO RAS). ( z = # 1 , where # is the coecient of
vertical attenuation of downwelling
ux of solar radi- ecosystems both in individual ocean regions [Campbell
ation.) In these missions many abiotic ecosystem pa-
rameters were evaluated with determinations of dimen-
sional structure, biomass, and production of picoplank-
ton, nanoplankton, and microphytoplankton, as well
as basic heterotrophic elements of communities such as
bacteria, protozoa, and various dimensional and trophic
of groups of mesoplankton and macroplankton. It was
shown that there exist close relationships between the
content of surface phytopigments and the above ecosys-
Copyright 96 by the American Geophysical Union.
0001{4370/96/3505{0009$18.00/1
643
seasons) and in actual (calendar seasons) time.
It should be noted that dierent authors have already
used satellite data to investigate seasonal changes in
and Aarup, 1992; Comiso et al., 1993; Muller-Korger et
al., 1989] and in the World Ocean as a whole [Banse and
English, 1994]. However, their analyses were limited to
examination of variability patterns of chlorophyll con-
centration in the surface layer of the ocean. In contrast,
we see the problem in evaluation of a seasonal variabil-
ity of stocks and in consideration of other components
of communities in the productive layer such as biomass
and production of phytoplankton, bacteria, and zoo-
plankton.
Attempts to evaluate stocks of diverse components
of epipelagic communities of dierent productive-
644 SHUSHKINA ET AL.: EPIPELAGIC ECOSYSTEMS OF PACIFIC OCEAN
 SHUSHKINA ET AL.: EPIPELAGIC ECOSYSTEMS OF PACIFIC OCEAN 645
geographical zones of the Pacic Ocean were under- developed program. The stock estimates were obtained
taken by us earlier [Vinogradov and Shushkina, 1988; through multiplying mean biomass of one or another
Shushkina and Vinogradov, 1988]. However, in the ab- component in waters of each surface chlorophyll grada-
sence of satellite data, the mapping of such zones could tion (see Table 1) by the area of waters featuring this
be conducted only provisionally, and we could allocate gradation during a specic season (see Table 2).
only three classes of waters, i.e., eutrophic, oligotrophic, We must note that the NASA algorithm yields quite
and intermediate classes, but it appeared unrealistic to satisfactory conformity to ground-truth chlorophyll
separate the seasonal variability in their congurations
and areas.
The satellite data on surface chlorophyll concentra-
tion represent a reliable basis for zoning the ocean in
productivity for dierent seasons, while the data set of
ground-truth ecosystem observations, available at the
IO RAS, can serve as a basis for evaluating average con-
centration and stocks of major elements of communities
in separate regions of temperate and tropical areas.
Materials and Methods
evaluations at tropical and moderate latitudes, while
in high latitudes and especially in coastal and terrige-
nous suspension rich regions this conformity is upset.
Thus as follows from the work by Sallivan et al. [1993],
satellite evaluations are twice as high as direct deter-
minations in the Southern Ocean. This ratio is even
higher in epicontinental arctic seas [Vinogradov et al.,
1994]. Therefore use of the area of regions of dierent
trophicity for calculations should hereinafter be slightly
adjusted.
The calculations of areas occupied by waters of di-
verse coloration were made for various climatic seasons
within the contours formed by boundaries of the Pa-
Information on biological parameters of epipelagic cic Ocean for circumpolar and temperate regions north
communities was acquired with the same technique in of 40N and south of 40S and for the tropical and
several ecosystem eld studies. They were conducted subtropical zones between 40N and 40S. The to-
on board the vessels of the P. P. Shirshov Institute of
Oceanology in 1968{1990 and continued in the Okhotsk
and Bering Seas in 1991{1993 in expeditions run by
VNIRO. The data from about 250 stations have been
analytically treated. A map of station positions was
presented by Vinogradov and Shushkina [1988]. Typi-
cal values of the aforementioned biological characteris-
tics, computed as geometric averages, were determined
for waters of each of the seven selected levels of chloro-
phyll concentration [Vinogradov et al., 1992] separately
for cold-temperate waters and for tropical and subtrop-
ical waters. The technique for collecting and process-
ing these data was reported earlier [Vinogradov, 1980;
Vinogradov et al., 1984; Vinogradov, 1983]. All obser-
vations were conducted in the daytime, and the mass of
organisms was determined through counting and mea-
suring them under a microscope and subsequent recal-
culation of biomass from dierent regression equations
and empirical formulas.
The relation between the concentration of chloro-
phyll a in the surface layer and biomass values is not
functional in character; therefore the modal values of
biomass were used as the most probable estimates of the
latter for each color gradation. The geometric means
were more adequate than the arithmetic means for the
goals to be attained. This provision was checked every
time for each parameter by estimating the arithmetic
and geometric means and the median. In the case of
biomass, the geometric mean was closer to the median
than the arithmetic mean.
The concentration of chlorophyll in satellite maps,
computed from the NASA algorithms, was presented in
seven gradations [Vinogradov et al., 1992], each of which
was associated with a certain color. An area of ocean
surface, occupied by waters of one of the color grada-
tal area inside the outline of the Pacic Ocean mea-
sured 172  106 km2 including 77  106 km2 for the
northern hemisphere and 95  106 km2 for the southern
hemisphere. A specially developed algorithm was em-
ployed to assign parameter values to spots of missing
surface chlorophyll data. When clouds covered major
parts of individual ocean regions in the map, as during
the antarctic climatic winter south of 60S, the ratio of
areas of dierent coloration was adopted to be the same
as in the adjacent cloudless regions.
The generalized primary data of satellite observations
were given to us by our American colleagues within the
framework of joint activity on the SeaWIFS program,
and we express our gratitude for this cooperation.
Results and Discussion
Table 1 presents the annual average values for biomass
of basic elements of epipelagic communities in regions
occupied by waters of dierent productivity (or col-
oration in satellite maps). It follows from Table 1,
that the phytoplankton biomass decreases from 1.4{
7.0 g C m 2 in high-productivity waters of three ini-
tial gradations to 0.4{0.3 g C m 2 in low-productivity
waters of 6{7 gradations, i.e., 5{20 times. The con-
centration of the heterotrophic microplankton (bacte-
ria and protozoa) diers to lesser degree (or approxi-
mately twofold). For the mesoplankton, the biomass
of both small sized and larger fractions varied from
eutrophic to oligotrophic waters sixfold to tenfold. It
should be noted that the small-sized mesoplankton ( Bm
( l < 3 mm)) comprised about one half of the total
mesoplankton biomass ( Bz ) in the moderately cold
water regions of high- and medium-productive waters
tions, was calculated for each of them with a specially (the rst to fourth gradations), while its contribution
646 SHUSHKINA ET AL.: EPIPELAGIC ECOSYSTEMS OF PACIFIC OCEAN

Table 2. Stocks of the major elements of communities within the upper 200-m layer of the Pacic Ocean during
diverse climatologic seasons (total for both hemispheres) [Bi ] = 106 t C
Phytoplankton, p Bacteria, b
Gradation Region Annual Spring Summer Fall Winter Annual
Spring Summer Fall Winter Average Average
1 P 21.5 16.4 22.4 8.8 17.3 6.3 6.4 6.4 2.6 5.4
T 10.0 6.4 6.4 9.3 8.0 1.2 0.8 0.8 1.2 1.0
2 P 6.3 5.5 7.1 4.0 5.7 1.8 1.6 2.1 1.2 1.7
T 3.4 4.9 3.5 4.2 4.0 0.6 0.3 0.8 0.7 0.6
3 P 15.8 15.5 13.2 14.6 14.8 8.2 8.0 6.8 7.6 7.6
T 10.9 5.6 12.9 19.6 12.2 3.4 1.7 4.0 6.1 3.8
4 P 14.7 12.0 5.5 16.2 12.1 9.8 8.0 3.7 10.8 8.1
T 10.2 5.7 9.5 14.7 10.0 7.4 4.1 6.9 10.7 7.3
5 P 5.7 7.7 3.7 8.7 6.4 3.5 4.7 2.3 5.3 4.0
T 5.1 4.9 6.9 7.1 6.0 5.9 5.6 7.9 8.1 6.9
6 P 2.2 3.2 5.4 3.0 3.4 4.0 5.8 9.9 5.4 6.3
T 15.9 18.4 20.0 18.2 18.1 14.5 16.8 18.1 16.6 16.5
7 T 19.0 20.1 14.1 9.5 15.7 19.9 21.0 14.7 10.0 16.4
Entire P 66.2 60.3 57.3 55.3 59.7 33.6 34.5 31.2 32.9 33.1
ocean P 141 126 131 138 13474.0 8652.9 8550.3 8453.2 8653.4 8652.5
T 74.5 66.0 73.3 82.6
Notes. P stands for polar and temperate regions and T stands for tropical and subtropical regions.
augmented to 70{80% in regions of lower productivity sults in nontrivial inferences. It appears true that these
(the fth to seventh gradation). The small-sized zoo- stocks in the ocean as a whole do not vary during a
plankton ( Bm ) comprised 70{85% of total zooplankton year. This takes place during both climatic seasons and
biomass in tropical regions for all gradations. calendar seasons, when the summer of one hemisphere
It is interesting to note well-dened distinctions in
the role played by predatory organisms in dierent geo-
graphical zones of the ocean. The predators contributed
4{8% of Bm into the small-sized mesoplankton in mod-
erately cold water regions, inhabited by cyclic commu-
nities. In contrast, their contribution increased to 15{
20% in the tropical regions, but in both cases this oc-
curred independently of water productivity. The preda-
tors comprise 12{18% of total mesoplankton biomass
corresponds to the winter of another and the equilib-
rium occurs at a level of (72{75)  106 t C in tropical
latitudes and at level of (54{69)  106 t C in temperate
latitudes (Table 2).
In tropical latitudes, phytoplankton stocks change
from 66  106 t C during climatic summer to
83  106 t C during climatic winter, and the opposite
occurs in temperate latitudes where stocks vary from
55  106 t C in winter to (66{60)  106 t C in spring
( Bz ) in highly productive waters (rst to third gra-
dations) of moderately cold water regions, but in the
tropical regions their contribution rises to 22{27%.
The conclusion about decrease in macroplankton
biomass ( Bq ) (organisms larger 3 cm) is less reliable,
since it is based on a smaller volume of materials ob-
tained with semiquantitative hauling tools. However,
the same tendency is also traceable in this case. After
and summer. It is as if the ocean breathes. The phyto-
plankton biomass grows from winter to spring and sum-
mer in relatively small eutrophic areas (gradations 1{3)
of temperate latitudes comprising about 15% of total
ocean area. However, the oligotrophic and ultraolig-
otrophic (gradations 6{7) aquatic areas, comprising on
an average of 55% of the ocean's surface, increase in
size during spring to summer. For example, in winter,
these introductory remarks on mean biomass values for
diverse elements of communities, consider their stocks
in ocean waters of dierent productivity. Especially
interesting is the intraannual stock variability during
various climatic seasons (Table 2), for example, during
the period of climatic summer, conterminous to cal-
endar summer in the northern hemisphere and to the
in regions of temperate zone of the rst to second gra-
dation, the phytoplankton stocks decrease 2{2.5 times,
but the extensive tropical ultraoligotrophic regions of
low phytoplankton biomass decrease twofold in area. As
this takes place, the total area of low-productive wa-
ters of sixth and seventh gradations contracts by about
30%, while the medium-productive regions of fourth
calendar winter in the southern hemisphere. and fth gradations, respectively, increase. Thus total
We begin with phytoplankton because it produces the phytoplankton stock in highly productive temperate
primary organic matter that occurs at the entrance of waters turns out to be smaller in biological winter than
the entire energy
ow into the oceanic ecosystems. The in spring or in summer, but, in contrast, it is larger in
analysis of seasonal changes in phytoplankton stocks re- poorer subtropical and tropical waters.
 SHUSHKINA ET AL.: EPIPELAGIC ECOSYSTEMS OF PACIFIC OCEAN 647

Table 2. (continued)
Protozoa, a Mesoplankton, z
Spring Summer Fall Winter Annual Spring Summer Fall Winter Annual
Average Average
1.8 1.4 1.8 0.7 1.4 38.4 29.3 38.7 15.7 30.5
0.4 0.2 0.2 0.3 0.3 2.4 1.5 1.5 2.3 1.9
0.6 0.5 0.7 0.4 0.6 9.4 8.2 10.6 6.0 8.6
0.2 0.3 0.2 0.3 0.3 1.4 2.0 1.4 1.7 1.6
2.0 1.9 1.6 1.8 1.8 38.7 38.1 32.3 35.8 36.2
1.1 0.6 1.2 1.9 1.2 8.1 4.1 9.6 14.5 9.1
2.8 2.3 1.0 3.1 2.3 32.6 26.8 12.2 36.0 26.9
2.3 1.3 2.1 3.3 2.3 17.4 9.6 16.1 25.0 17.0
0.7 0.9 0.4 1.0 0.8 4.2 5.6 2.7 6.3 4.7
1.8 1.7 2.4 2.5 2.1 10.4 9.9 14.0 14.4 12.2
0.7 1.0 1.6 0.9 1.0 2.5 3.6 6.2 3.4 3.9
6.8 7.9 8.5 7.8 7.7 19.2 22.2 24.0 22.0 21.8
5.0 5.3 3.7 2.5 4.2 12.2 12.9 9.0 6.1 10.1
8.0 8.0 7.1 7.9 7.9 125.8 111.6 102.7 103.2 110.8
17.6 17.3 18.3 18.6 18.1 71.1 62.2 75.6 86.0 73.7
26 25 25 27 26 197 174 178 189 184

As a result, the mean quantity of phytoplankton re- ina, 1988], especially for medium-productive (the fourth
mains at about the same level for the entire aquatic and fth gradations) and oligotrophic (the sixth grada-
area of the Pacic Ocean, ranging from 126  106 t C
to 141  106 t C in dierent seasons, with an annual
average of 134  106 t C (Table 2).
It is necessary to point out that the use of satellite
information for distinguishing regions of dierent pro-
ductivity resulted in the correction of an area of highly
productive regions toward its reduction in reference to
tion) waters that occupy 65% the area of the Pacic
Ocean. At the same time, a correction of highly pro-
ductive areas occurred in favor of reduced values [Vino-
gradov et al., 1995]. As a result, the estimate for the
total annual primary production turned out to be close
to the preceding estimates [Vinogradov and Shushkina,
1988]. The latter computed from geometric means was
earlier eld observations [Vinogradov and Shushkina, as high as  21  109 t C yr 1 , while the estimate from
1988]; this especially concerns the hypertrophic waters satellite data equals to 27  109 t C yr 1 . If the cor-
of the rst gradation. In addition, new eld data on rections for the so-called
ask eect are made according
biomass were obtained that re
ect periods of phyto- to Shushkina et al. [1987], this value becomes close to
plankton blooms in dierent regions. Also, careful anal- 46  109 t C yr 1 for the whole of the Pacic Ocean,
ysis of all eld and experimental materials at our dis-
posal have been performed when only those data were
examined that were obtained with techniques enabling
one to account for microplankton, nanoplankton, and
picoplankton. Finally, it is important that it has be-
come possible to use realistic information on individ-
ual seasons, while earlier we were forced to deal mostly
with summer season materials and to take into account
the duration of the vegetation period for circumpolar
and temperate regions. This resulted in a substantial
almost twofold decrease in the estimate of average an-
nual stock of phytoplankton as compared to an earlier
reported value [Shushkina and Vinogradov, 1988]. In
recent years, advances in techniques for determination
uctuations of their amount are less pronounced. For
while in the work by Vinogradov and Shushkina [1988]
it was estimated as 36  109 t C yr 1 .
Bacteria and Protozoa (Heterotrophic
Microplankton)
While the distribution of phytoplankton in the ocean
is rigorously subjected to biological latitudinal and cir-
cumcontinental zonality [Bogorov and Zenkevich, 1966;
Vinogradov and Shushkina, 1989; Zenkevich, 1948;
Hentschel, 1936] that reveals the richness of coastal and
high-latitude areas and the poverty of tropical anticy-
clonic gyres, the stocks of heterotrophic microplank-
ton are more evenly distributed and the patterns of
of primary production in medium- and low-productive these reasons, the uniform eld of gures in Table 2 is
ocean regions have yielded new estimates [Vedernikov not odd. It is necessary only to pay attention to the
and Konovalov, 1990] that turned out to be about twice higher biomass of bacteria and protozoa in the low-
as high as was earlier reported [Vinogradov and Shushk- production tropical regions of fth to sixth gradations
648 SHUSHKINA ET AL.: EPIPELAGIC ECOSYSTEMS OF PACIFIC OCEAN

Table 3. Distribution of annual average stocks of major highly productive and low-production regions, respec-
components of epipelagic communities over regions of tively.
diverse trophicity (in percents) As a result, almost one half of the total zooplank-
Phyto- Bacte- Proto- Meso- ton stock (48%) is concentrated in productive regions
Gra- Water of the rst to third gradations, occupying about 15%
dation trophicity plank- ton ria zoa plank-
ton ocean's area, and almost the same fraction (52%) occurs
in poorer mesotrophic and oligotrophic waters. The
1 Hypertrophic 19 7 6 18 same ratios are characteristic of phytoplankton. For
2{3 Eutrophic 27 16 15 30 bacteria and protozoa, an inverse pattern takes place:
4{5 Mesotrophic 26 31 29 33 the mesotrophic and oligotrophic waters comprise about
6{7 Oligotrophic 28 46 50 19 80% of stocks (Table 3).
Total 100 100 100 100 Despite the longer life cycles of mesoplankton, its
mean annual stock in the ocean (184  106 t C) turns
out to be smaller than the supply of the primary
relative to the temperate ones, as was already reported food, i.e., of phytoplankton, bacteria, and the proto-
many times, and hence to the more important contri- zoa, which measures 246  106 t C (Table 2).
bution of tropical regions to the total mass of micro- Comparing the obtained mean stock of zooplankton
heterotrophs in the ocean (on the average, 53  106 t C to the earlier reported estimates [Shushkina and Vino-
against 33  106 t C for bacteria, and 18  106 t C gradov, 1988] shows that its value remained virtually the
against 8  106 t C for protozoa). The total stock same, or (185{189)  106 t C, in distinction to the mi-
of bacteria in the ocean layer from 0 to 200 m mea- croplankton stocks (the primary food). Probably, this
sures 86  106 t C, while that of protozoa equals to is due to greater accuracy of maps for zooplankton dis-
26  106 t C, whose sum is close to the phytoplankton tributions in reference to phytoplankton maps that we
stocks. employed in the preceding study [Shushkina and Vino-
In determination of bacteria and protozoa, only those gradov, 1988].
data were taken into account that were obtained with Once again, seasonal dierences for the ocean as a
the help of
uorescent microscopy and
uorochrome whole appear to be insignicant: mesoplankton stocks
pigments. As in case of phytoplankton, the computed
values for stocks of bacteria and protozoa (Table 2)
turned out to be about 1.5{2 times lower than the
earlier reported estimates [Shushkina and Vinogradov,
1988].
Mesoplankton
Similar to phytoplankton distribution, mesoplankton
for the entire ocean in the upper 200-m layer vary from
197106 t C to 174106 t C, diering only by 6% of its
mean value. Certainly, it should be taken into account
that in autumn and winter the major populations of the
upper interzonal species descend from the 0 to 200-m
layer at greater depths in moderately cold water regions
inhabited by cyclical communities. However, in spring
and summer, when their populations are still feeding in
the subsurface layer, total mesoplankton stock increases
distribution is subjected to patterns of latitudinal and
circumcontinental zonality. Also, due to the longer life
cycle of zooplankton animals in comparison to that of
phytoplankton, the values of biomass occurring during
a year are closer to annual stocks than occurs in the case
uence) are relatively modest owing to the
insignicantly.
We note again that total stocks of mesozooplank-
ton in eutrophic waters of tropical latitudes (mostly,
the aquatic areas of upwelling and of regions subjected
to their in
of phytoplankton. This is particularly true for temper-
ate latitudes. The amount of zooplankton in cyclical
communities of temperate latitudes, where life cycles of
dominating interzonal species feature annual or nearly
annual duration, proves to be larger than in highly bal-
anced communities of tropical oligotrophic regions (Ta-
ble 2).
For this reason, 40% of the total mass of mesozoo-
plankton is concentrated in hypertrophic and eutrophic
insignicance of the areas occupied by these waters.
Moreover, in these waters populated by cyclic communi-
ties where a major portion of mesoplankton feeds during
short periods of rapid phytoplankton growth and then
descends below the 200-m level as upwelling relaxes, the
average zooplankton biomass in the upper 200-m layer
turns out to be lower than in productive waters of tem-
perate latitudes (Table 1). This alone explains the fact
that the upwelling regions play a secondary role in for-
waters (the rst to third gradations) of polar regions mation of zooplankton stocks, while they play a leading
comprising 11% of the ocean area, while only 8% of role in forming phytoplankton stocks in the productive
these animals inhabit tropical waters of the same gra-
dations. In low-production waters of the sixth and sev-
enth gradations the pattern is inverted: polar waters
comprise about 2%, while tropical water contains 17%
of the annual average stock of zooplankton of the entire
ocean. In these relationships, the medium-production
ocean areas. In the high-production regions of the rst
to third gradations, the polar zones and tropical up-
welling regions yield on the average 28% and 18% of
total phytoplankton stocks, respectively, whereas cor-
responding contributions into the zooplankton stocks
are 41% and 7%.
waters of fourth and fth gradations are closer to the The total zooplankton stocks are large in tropical
 SHUSHKINA ET AL.: EPIPELAGIC ECOSYSTEMS OF PACIFIC OCEAN 649
oligotrophic regions owing to their enormous area. Muller-Korger, F. E., C. R. McClain, and T. R. Fisher, et
These stocks are close to contribution from temperate al., Pigment distribution in the Caribbean Sea: Observa-
latitudes (40% and 60%, respectively). If one considers tion from space, Prog. Oceanogr., 23, 23{64, 1989.
the calendar seasons when a decrease in areas in one Shushkina, E. A., and M. E. Vinogradov, Quantitative char-
hemisphere is compensated by an increase in another
one, the value of total stock does not vary. It is an-
other thing to consider dierent climatic seasons for
the ocean as a whole. In this case, total zooplankton
stocks prove to be twice as low as in summer owing
asks during determination of primary and bacterial pro-
to winter decrease in areas of ultraoligotrophic gyres
(seventh gradation). These
uctuations in oligotrophic Sullivan, C. W., K. R. Arrigo, and C. R. McClain, et al., Dis-
acterization of pelagic population of the Pacic Ocean,
Plankton biomass and destruction-production processes,
Oceanology, Engl. Transl., 28, No. 6, 992{1000, 1988.
Shushkina, E. A., M. E. Vinogradov, and B. V. Konovalov,
et al., Changes in microplankton community in testing
duction, Izv. Acad. Sci. USSR Biologiya, 1, 42{54, 1987.
tribution of phytoplankton blooms in the Southern Ocean,
regions of the sixth gradation are less signicant. Science, 262, 1832{1837, 1989.
While considering seasonal changes in phytoplankton Vedernikov, V. I., and B. V. Konovalov, Primary production
stocks, it was found that generally they occur in an-
tiphase in temperate and tropical zones due to alter-
ation of their areas featuring distinct productivity lev-
els. Total zooplankton stock in the 0 to 200-m layer also
remains almost constant during the year, varying from
174  106 t C to 197  106 t C. A similar conclusion
can be drawn concerning the total plankton biomass
B0 = Bp + Bb + Ba + Bz , which measures 450  106 t C
in spring, 410  106 t C in summer, 418  106 t C in au-
tumn, and 440  106 t C in winter at an annual average
and chlorophyll in the northern Pacic Ocean in July-
September 1990, Trudy Instituta Okeanologii RAN, 131,
16{31, 1994.
Vinogradov, M. E., Distribution of trophic groups of
zooplankton in cross section o Pacasmayo Peninsula
(70 370 S), in Pelagic Ecosystems of the Peruvian Region,
pp. 159{166, Nauka, Moscow, 1980.
Vinogradov, M. E. (Ed.), Modern Techniques for Quan-
titative Evaluation of Distribution of Marine Plankton,
279 pp., Nauka, Moscow, 1983.
Vinogradov, M. E., and E. A. Shushkina, Quantitative char-
of 430  106 t C.
Apparently, the annual stability of stocks of both
autotrophic and heterotrophic plankton, and, subse-
quently, of metabolism processes in the ocean has far-
reaching global consequences related to special features
acterization of the pelagic population of the Pacic Ocean,
Productive regions and estimates of the primary produc-
tion of photosynthesis, Oceanology, Engl. Transl., 28,
No. 5, 819{827, 1988.
Vinogradov, M. E., and E. A. Shushkina, The macroscale
distribution of quantitative characteristics of plankton in
of the carbon cycle, distributions of CO2 and O2 , and the Pacic Ocean, Oceanology, Engl. Transl., 29, No. 1,
to other biospheric parameters. 121{126, 1989.
Vinogradov, M. E., E. A. Shushkina, G. N. Arnautov, and
Acknowledgments. This work was supported by the E. I. Musaeva, Comparative characterization of meso-
Russian Fund for Basic Research (project 94{05{17405), by plankton biomass in waters of diverse trophicity in the
the Ministry of Science Program \Global changes of envi- southwestern Pacic Ocean, in Frontal Zones of the South-
ronment(Wednesday) both climate," and by the ISF grant
Supplementary Grants Program N SAYOOO under the pro-
gram SeaWIFS.
ux in
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