ContinentalShelfResearch,Vol. 10, No. 4, pp. 355-367, 1990. 0278--4343/90$3.00 + 0.

00
Printed in Great Britain. ~) 1990PergamonPressplc

Sulfate reduction in shelf sediments in the upwelling region off

Central Peru

HENRIK FOSSING*

(Received 12 June 1989; accepted 8 February 1990)

Abstract---Concentration profiles of sulfate, hydrogen sulfide, ferrous sulfide, elemental sulfur,

and pyrite are presented from five stations from the upwelling region off Peru near 15°S together
with profiles of water content, density and organic content. Four of the stations were located within
the oxygen minimum zone (80-500 m depth) and one was located at 2650 m depth. Sulfate
reduction rates were determined from radiotracer measurements on all stations by acid distillation
followed by chromium reduction. Reduced 35S recovered as acid volatile sulfide (H2S and FeS) was
compared to 35S recovered as chromium reducible sulfur (S ° and FeS2). In the top 10 cm of the
sediment 15-40% of 35Sred was recovered as acid volatile sulfide, deeper into the sediment it was
less than 10%. Maximum reduction rates of 68-312 nmol cm -3 day- 1 were observed within the top
4 cm of sediment from stations in the oxygen minimum zone. Integrating sulfate reduction rates
over whole sediment cores under the oxygen depleted water column gave an average rate of 20 +
l l m m o l SO 2- m -2 day - l (n = 3). This corresponds to the mineralization of 9-29% of the
planktonic primary production in the overlying water column,

INTRODUCTION

THE upwelling region off Peru has very high primary productivities of 1-10 g C m -2 day-1
(RrmER et al., 1971; LEE and CsoNI1q, 1982; HErqFaCrlSand FAsmN~rOrq, 1984). This is 10
times higher than the average primary productivity of the ocean, 0.1-0.2 g C m -2 day -1
(MENzEL and RrraEt, 1960). Up to 90% of the primary production is remineralized within
the water column (LEE and CRol~Irq, 1982; RowE, 1985), resulting in oxygen depletion
from 80 to 500 m depth (<0.1 ml 021-1) in the upweUing zone.
When oxygen is depleted the microbial degradation of organic material takes place with
NO3 or NO2 as terminal electron acceptors (CoDISPOTI, 1983; CODISPOTIand CnPaSTEN-
SEN, 1985; CoI)ISPOrIet al., 1986). Total depletion of NO3 in the water column has been
observed only once in the Peru upwelling area when H2S was detected (DUGDALEet al.,
1977). In the sediment the mineralization processes proceed with NO3, NO2 and SO2- as
electron acceptors. Despite the importance of sulfate as the terminal electron acceptor in
marine sediments in general (FRoELICHet al., 1979; JORGENSEN,1983; LEIN, 1984; Sr~'lUr~G,
1987) only one study of sulfate reduction has been reported in sediments influenced by
upwelling (Rowe and HOWARTIa,1985). In the study of Rowe and Howarth, maximum
sulfate reduction rates of approx. 30 nmol cm -3 day- 1 were measured between 9 and 12 cm
depth and total sulfate reduction rates were 5--6 mmol m -2 day -i, calculated by

*Institute of Ecology and Genetics, University of Aarhus, Ny Munkegade, DK-8000 Aarhus C, Denmark.

355
356 H. FOSSING

14°50 S' ~
~~-~:":!:. ,t/ R/V Moana Wave
:~'aca
, • July 1987
5.:..:' '~
~ i~.....;,,.i:...:.
" 2~... -

15° 10 00, ! ~ : ~ ~i!.-.-:.....

\ ~ x ~
ni:a San Nicolas
.w.::......

• 10 c~0tm ZN~TB~
0. 5 10 t,s 2.0
KILOMETERS
].5o30 f I i
76°00 'W 75°30 ' 75°00 '
Fig. 1. Sampling locations in the upwelling region off Central Peru.

integration over the depth of the cores. These rates were 10 to 100-fold higher than
observed ocean-wide at comparable water depth (SrYPdNG, 1987).
Presented here are sulfate reduction rate measurements from five stations visited during
PUBS I expedition with R.V. Moana Wave (cruise 87-08), July 1987. The purpose is to
report how fractions of reduced sulfate are incorporated into acid volatile sulfide (AVS:
H2S and FeS) and chromium reducible sulfur (CRS: S° and FeS2). In addition, the
sediment chemistry of SO42-, H2S, S°, FeS and FeS2 are also reported and discussed.

MATERIAL AND METHODS

The depths ranged from 130 to 2500 m and ranged in bottom water oxygen content from
<2 to 100% of air saturation. Stations 4, 6, 8 and 10 were located 15-450 km from the coast
near 15°S, whereas Sta. 2 was positioned further north at 11°S (Fig. 1, Table 1).

Sampling and sediment handling

Sediment was obtained from all stations by a box corer (BC) (Sandia-Hessler MK-III
Ocean Instruments, San Diego; 0.25 m 2, depth 30 cm). Sediment cores of varying lengths
were also sampled with gravity corer (GC) (200 cm length, 15 cm diameter) at Stas 2, 4 and
8 and at Sta. 6 with a piston corer (PC) (500 cm length, 10 cm diameter). Sediment was also
sampled by a Soutar corer (SC) (0.1 m 2, depth 1 m) at Sta. 4 (Table 1).
Sediments from BC and SC were visually inspected for an undisturbed surface. The GC
and PC were always recovered with the surface layer lost. The loss of sediment was
estimated from the chemical data when these observations were aligned with the data
obtained from the BC. Subsampling was done with Plexiglas coring tubes pressed into the
sediment. To avoid compression during subsampling, a piston mounted inside the coring
tube was pulled simultaneously as the tube was pressed into the mud. Subcores of 15-20 cm
length were sampled in triplicate from BC, GC, PC and SC with 3 cm diameter coring
 Sulfate reduction in shelf sediments 357

Table 1. Sampling stations and sediment cores sampled

Water Bottom Oxygen Top

depth temp. content Core lostt
Station Location (m) (°C) (ml 021 -l) type* (cm)

2 11°04.2'S 255 14 0.07 BC 0

78°31.2'W GC 11
4 15°06.2'S 268 14 0.04 BC 6
75°42.1 'W SC 0
GC 19
6 15°11.5'S 502 15 0.12 BC 0
75°34.5'W PC 8
8 15°00.0'S 135 16 0.08 BC 0
75°39.2'W GC 16
10 15°20'S 2650 6 Saturation BC 0
75°50'W

*BC: MK-III box corer; GC: large diameter gravity corer; SC: Soutar type box corer; PC: piston corer.
tThe calculated thickness of sediment that was blown off from the surface when sampled.

tubes. The gravity and piston cores were pushed from the coreliners as consecutive
subcores were taken. The subcores were all sealed with butyl rubber stoppers and stored
by in situ t e m p e r a t u r e <1.5 h before radiolabeling.

Analyses
Sulfate reduction measurement. Injections of 2 ktl carrier-free 35SO2- (200 MBq m1-1)
( A m e r s h a m Corp.) were done horizontally, at 1 cm intervals, into the sediment through
silicone-stoppered ports (JORGENSEN, 1978). The radiolabeled sediment was incubated for
8-16 h at in situ t e m p e r a t u r e (cf. Table 1) in a thermostated van. To prevent AVS
oxidation, the cores were extruded in 1 or 2 cm steps from the coreliner, immediately cut
into 10 ml 5% zinc acetate fixative in a 20 ml vol. plastic vial, thoroughly mixed, and
weighed. The sediment was then centrifuged within the plastic vial and radioactive sulfate
determined from 5 ml of a diluted subsample of the supernatant mixed with 5 ml of
scintillation liquid (Dynagel, B a k e r Chemicals).
The fraction of 35SO2- reduced during incubation was determined from an acid
distillation followed by a chromium reduction (ZHABINAand Volkov, 1978; WESTRICH,
1983; HOWARTH and JORGENSEN, 1984; CANFIELD et al., 1986, FOSSING and JORGENSEN,
1989). The sediment pellet was washed twice and homogenized. A 2-3 g subsample was
transferred to a reaction flask with condenser and 12 N HCI was added to a final
concentration of 2 N HCI. Distillation c o m m e n c e d for 30 miD and liberated all A V S as
H2S, which was collected as ZnS in two sequential traps containing 10 ml of 5% zinc
acetate (ZnAc) plus a drop of antifoam. The traps were then renewed and chromium
reduction proceeded on the same sediment with Cr 2+ and HC1 added. All CRS was
dissolved as H2S and carried to the traps within 40 miD. For both digestions, the pairs of
Zn35S traps were pooled, a 5 ml subsample was mixed with 5 ml of scintillation fluid, and
35S was counted on a scintillation counter (Packard, Tri-Carb 2200 CA). The sulfate
358 H. FOSSlN~

reduction rates were calculated as described by JORGENSEN (1978) and FOSSlNG and
JOR~ENSEN (1989).
Radiolabeled elemental sulfur was extracted with 5 ml of CS2 from 2 g of the washed and
homogenized sediment. A 2 ml subsample of the CS2 was vaporized, the 35S° was
redissolved into 5 ml of scintillation liquid, and counted after addition of 5 ml distilled
H20. All radioactivities were quench corrected.
Chemical analyses. Sulfate and free sulfide were obtained from porewater by pressure
filtering through a 0.45/~m membrane filter (Millipore) under a gas-impermeable latex
membrane. The first 5 drops of porewater were discharged. To minimize exposure to the
atmosphere up to 2 ml of porewater (determined by weight) was transferred directly into
1.0 ml of 2% ZnCI 2 fixative in a plastic vial. The concentration of sulfate was determined
on diluted porewater samples by non-suppressed ion chromatography (Waters) using an
anion exchange column and conductivity detector. Free sulfide was measured spectro-
photometrically from precipitated ZnS as described by CI~INE (1969), as were concen-
trations of AVS and CRS on subsamples of the ZnS from the H235Sdistillations. Elemental
sulfur concentration was measured on a 1 ml subsample of CS2 from the S° extraction as
described by TROELSENand JORGENSEN(1982).
Porosity and organic content. Known volumes of sediment were transferred into 20 ml
plastic vials and weighed. The densities were determined and used to recalculate rates and
concentrations from weight to volume. The sediment was then dried overnight at 105°C to
estimate the water content. Porosity was calculated from density and % water content
(w/w). Organic content was measured as loss on ignition from the dried sediment (4 h at
450°C).

RESULTS
The sediment cores were all sampled from the oxygen minimum zone (<0.1 ml 021-1)
except from the deep Sta. 10 at 2650 m, where the oxygen concentration was estimated to
be at air saturation. From each station, except Sta. 10, the profiles of water content,
density, and AVS in the sediments obtained by box, gravity, piston, or soutar coring were
aligned in order to estimate how much of the surface was lost when the sediment was
sampled (Table 1). All graphic presentations are based on these depth estimates.
In the oxygen minimum zone the upper 5-10 cm of the sediment was brown to blackish
green, with a gelatin-like consistency and contained fish scales, bones, and other organic
debris (in the top 1 cm of fresh sediment, 150-250 g organic material m-2). The porosity
ranged from 95 to >99% in the top 10 cm. At Stas 2, 4 and 8 (Figs 2, 3 and 5), filamentous
colorless sulfur bacteria of the genus Thioploca were also observed within the upper 15 cm
of sediment (GALLARDO,1977), with the highest density at Sta. 8. Deeper into the core the
sediment changed to an olive green diatomaceous ooze with yellow laminations and the
water content decreased. At Sta. 6 a compact layer of mud was located from 10 to 20 cm on
top of sediment with a higher porosity (Fig. 4).

Sulfate
At all stations the sulfate concentration was approx. 28 mM in the top 10-20 cm of the
cores (Figs 2-6). Below this depth the concentration decreased. Only at Sta. 4 did SOZ4-
become depleted within the length of the core, at 140 cm depth.
 Sulfate reduction in shelf sediments 359

WATER {ml "9-11 DENSITY (g .cm-al x ORGANIC (w/w) 504~- (mM) HzS (mM)
0.6 0 .B 1.0 1.5 2 0 20 40 10 20 30 5 10
i , i ~

°~w
i
g-
t %°
-~ 50.
eo

loo qje
41 ee

t5C
SRRAvs SRRAvs÷cRS
S° (~mol .cm-3) FeS {#tool"cm-3) FeS~ (~mol S .cm-3) nm01-cm-~.day -t rim01 "cm- 3 -(lay - i
0.1 0.2 0 0.5 ! 50 I00 150 0 5 I0 50 100

• ~O0

50. i

/
/
1oo

150.

Fig. 2. Station2. Porewaterand solid phase datawith depth in core. Organiccontent determined
as percent loss on ignition from dried sediment. SRRAvs: sulfate reduction rate determined from
acid distillation of H2S + FeS. SRRAvs+cRS: sulfate reduction rate determined from acid
distillation followed by chromiumreduction of S° + FeS2. Increase in sulfate reduction rate below
40 cm (broken line) may be an artifact.

Free sulfide
N o free sulfide was detected within the upper 10-15 cm at any of the stations
investigated, but below that depth free sulfide concentration increased. This increase was
reversely related to the sulfate concentration and reached the highest concentration, 15
mM, at Sta. 4.

Ferrous sulfide
The concentration of FeS was calculated from the difference between A V S and free
sulfide concentrations. In the oxygen minimum zone, FeS was detected between the
sediment surface and the boundary of the free sulfides, which gave the sediment a black
appearance. The maximum concentrations of FeS varied from 0.5/zmol cm -3 at Sta. 2 to
2 . 0 / z m o l cm -3 at Sta. 8, with maxima at a depth of 11 and 0.5 cm, respectively. As
expected from the grey-colored (oxidized) top sediment at Sta. 10, FeS was first detected
deeper in the sediment where the concentration increased up to 5/zmol c m - 3 at 10 cm.

Elemental sulfur
Elemental sulfur was not detected at Sta. 4 and only at a very low concentration ( < 0 . 1
/zmol cm -a) within the upper 5 cm at Stas 2 and 8. A maximum concentration of 1.57/zmol
360 H. FOSSING

S ° c m - 3 was measured at Sta. 6 at a depth of 3.5 cm, but S ° was depleted at 10 cm. The more
oxidized sediment of Sta. 10 showed an increasing concentration of elemental sulfur below
5 cm.

Pyrite
Pyrite normally exceeds elemental sulfur by orders of magnitudes in marine sediments
(GOLDHABER and KAPLAN, 1974) and therefore the bulk concentration of CRS should be
FeS2. Pyrite was the largest reduced sulfur pool in the sediment at all stations but low in
concentration near the surface (Figs 2-6).

Sulfate reduction rates

Sulfate reduction rates (SRR) based on A V S alone (SRRAvs) were significantly lower
than rates determined from the combined distillation of A V S and CRS (Figs 2--6). When
most extreme, SRRAvs+cR s exceeded SRRAvs by more than 10-fold as in the top
sediment of Sta. 2. Sulfate reduction was most intense a few cm below the sediment surface
on all stations in the oxygen minimum zone. The reduction rate decreased rapidly with
depth except at Sta. 2 where sulfate reduction increased again below 40 cm (broken line in
Fig. 2). I cannot explain this SRR increase as it is not correlated to any decrease in sulfate
concentration. At present, thus, I consider this observation to be an artifact. Station 10
(Fig. 6) revealed no sulfate reduction in the upper 5 cm based on A V 3 5 S alone, but CR35S
indicated that sulfate reduction was taking place in the oxidized part of the sediment.

WATER (ml-g-tl OENSITY (g.cm-3) ORGANIC (w/w) S04z- (mM) HzS (mM)
o6 o8 Ip 1.5
i
2
a
20 40,
10
i
20 30
i , i
10 20 30

,l k
-~ 50

C3 ioo
oO
•e
15C
SRRAvs SRRAvs÷CRS
S = (#tool-cm-3l FeS (amol cm-31 FeSz (t/tool S .cm-3) nmol .cm - ~ .day - ~ nmQ1 .cm - 3 - d a y - ~
0.1 0.2 I 2 0 50 100 1500 10 20 30 0 50 100
i i _ , i i ~I~ i i i

• J~ J%
i
-~ 50~
u

~oo

i50

Fig. 3. Station 4. Porewater and solid phase data with depth in core. See Fig. 2 legend.
 Sulfate reduction in shelf sediments 361

WATER (ml .g-q DENSITY (g "cm-3] % ORGANIC [w/w) SO4a- (raM) HaS (mM)
. 0.6 0.8 1.0 t.5 2 0 20 40 O 1o 2o 30 o 10 20 30
i , a ... n ...... .. , i i i
q~ raP"-
e

100
E • • •
• -

200. • o• °o NA
••

300. .2
%

,100. o°

SRRAvs SRRAvs+cRs
S ° (umol-cm-3) AVS (#tool .cm-3) FeSa (#mol S .cm-3) n m o l .cm - a . d a y - ~ lmol,cm-3.day -~

Oq
•
•
1
w J~ '.
2
i
2.5 5 200 ,u,0
,L.v' ,~,. ~ -
50
'
tOO 200 400

t
tO0
• 50 50

200.

tD0 tO0.
300. • o
• 3DD- 3001

400. • 400 400

Fig. 4. Station 6. Porewater and solid phase data with depth in core. See Fig. 2 legend. NA: not
analysed.

WATER [mlg -a} DENSITY [g 'cm-3} % ORGANIC (w/wl SO4z- (raM) HzS lmMl
0.6 0.8 10

;,':.
1.0 l.fl 2 0 20 40 20 30 5 1o
i , i ~ u . . . . . . . . . . , n i . . . . . . . . , , u

$ a i 't
20 | e°

It
t~ 40

60- •

B0-

100
SRRAvs SRRAvs+cRs
S° (umol 'cm-3} FeS (#mol "CFF3) FeSa [#tool S .cm-3) nmol .cm-3 -day-~ nmol -cm-3 .clay-~
o.o 0.1 0.2 2 0
g ,
50
i
100
i
150
g
10 20 30 0 |00 200
Ok • ,f n ' 'e~ I,
P
20 @ ,,,o • •

E 6o~

80
t
1(10

Fig. 5. Station 8. Porewater and solid phase data with depth in core. See Fig. 2 legend.
362 H. Foss]No

WATER [ml .g-~} DENSITY[g .cm-al x ORGANIC {w/w) SO4z - [mM) HaS (mM}
0,6 . 0.r.
, * t.,0 . . . .
t.5 , . . .
2 0
,
20\ 40 1
i
tO 20 30 0 tp 2p 3p
", :"
• , . , , oo

5-
E •. •

t0-
,.=,
o
15.

20
SRRAvs SRRAvs+CRS
S= (/Jmol .cm-3) FeS (umol "cm-31 FeSa [/Jmol S .cm-3) nmol .cm-3 .aay-= nmol.cm-=.day-~
u.,1 0.2 0 2.5 5I 0 25 50 0 50 tOOl 50 tO0
8 I I I I I I 'p . . . . . . . . . . . . . . . . . . , , i

• e•

E
5.
- . .,=
t0,

t5

2C

Fig. 6. Station 10. Porewater and solid phase data with depth in core. See Fig. 2 legend.

DISCUSSION

More than 50% of the total sulfate reduction took place within the upper 20 cm of the
sediment at Stas 4, 6 and 8 (Table 2). At these stations, the SRR reached a maximum 1-4
cm below the sediment surface (Figs 3-5). Oxygen concentrations were measured with
microelectrodes and showed that oxygen decreased from <0.1 ml 02 1-1 (<5 #M 02) at

Table 2. Sulfate reduction rate per m 2 o f the stations investigated. Sulfate

reduction rates integrated over the full core length and the top 20 cm o f the
sediment are compared to reduced 35S (35Sred) recovered in AVS in % o f
total 35Sred (AV35S + CR35S)

Sulfate reduction rate

(mmol SO 2- m -2 day -n)

Full core length 0-20 cm

Core 355re d 355re d

length in AVS in A V S
Station (cm) SRR (%) SRR (%)

2 100 32.5* 7.1 6.0 11.3

4 160 8.8 26.1 4.4 32.3
6 400 34.3 20.4 25.5 19.6
8 80 16.5 14.1 9.3 16.8
10 10 5.2 32.0 -- --

* Overestimated: see text.

 Sulfate reduction in shelfsediments 363

the surface to zero within <2 mm (NELSON and FOSSING, unpublished results). The oxygen
consumption was <5 mmol 02 m -2 day-1 as calculated from this oxygen gradient. Oxygen
was therefore only of minor importance for the mineralization of organic matter in the
sediment compared to sulfate. As nitrate and nitrite were not measured in the sediment on
this cruise it is not possible to estimate the importance of denitrification on these stations.

Sulfate reduction in sediments under the oxygen minimum zone

Sulfate which was reduced in the top 15 cm was immediately re-oxidized or precipitated
with ferrous iron, as no free sulfide was detected. In the sediment tops at Stas 2, 4 and 8,
colorless sulfur bacteria of the genus Thioploca are probably associated with the oxidation
of H2Sto elemental sulfur (LARKINand STROHL,1983; MAIERand GALLARDO,1984). So far,
the mechanism of this oxidation has not been elucidated, as Thioploca has never been
successfully cultured.
The relative recovery of reduced 35S (355red) in AVS showed an exponential decrease
with depth (Fig. 7). Between 15 and 40% of the sulfate reduced within the upper 10 cm of
the sediment was recovered as AVS. Less than 10% of 35Sred was recovered as AV35S
below 20 cm where a considerable sulfate reduction was still taking place (Table 2).
Significantly larger fractions of 355red have been recovered in AVS in shallower marine
sediments and salt marshes, typically 50-90% (WESTmCH, 1983; HOWARTHand JORGEN-
SEN, 1984; HOWES et al., 1984; KING et al., 1985; FOSSlNGand JORGENSEN, 1989). LEIN et al.
(1982) reported as little as 1% of reduced 35S in AvaSs during studies of sulfate reduction
in the Baltic Sea. The mechanism of tracer incorporation into the sulfur compounds that
comprise the CRS pool, mainly FeS2 is still not clear. The CRS pool comprises pyrite,
elemental sulfur, and other reduced sulfur compounds that are not dissolved during cold
acid distillation, e.g. greigite (BERNER, personal communication in HOWARTHand JORGEN-
SEN, 1984). Also elemental sulfur that is formed from oxidation of H2S by Fe 3÷ during acid
distillation (BERNER,1964, 1974) is recovered by the chromium reduction. However, in
nearshore sediments and in estuaries this fraction has been found only to vary from I to 5%
of the reduced 355 (HOWARTHand JORGENSEN, 1984; THODE-ANDERSEN and JORGENSEN,
1989).

100

o10
ta

ot

• , % **
1
t i i

0.1 1 10 100 1000

DEPTH (cm)
Fig. 7. Recoveryof reduced 35S (35S,~d)in AVS in percent of total reduced 3SS (=AV3SS +
CR35S) vs depth (cm) into the sediment. A double logarithmicplot of 115 data points from the
oxygenminimumzone (r = -0.67).
 Sulfatereductionin shelfsediments 365

they compared the measured SRR to rates estimated from SO~4- fluxes according to Fick's
first law of diffusion (cf. ROWEand HOWARTH,1985). However, they argued that the higher
SRR most likely were explained by an overestimation of the SO~4- flux at the interface
because they "did not know the distribution of SO42- in the bottom few centimeters of the
water column or within the top 3 cm of the sediment." (cited). To my concern, the
difference could also be explained from the low recovery of radiolabeled CRS. Higher
SRR than measured by Rowe and Howarth can be estimated from SO42- concentrations at
three stations in the same area, reported by HENRICHSand FARRINGTON(1984). Based on
these observations, I have estimated the SRR from the SO42- flux using Fick's first law of
diffusion and the same parameters as did Rowe and Howarth. For comparison these SRR
are presented in Table 3.
Based on SRR in this study, I find an average rate of 20 _+ 11 mmol SO ]- m -2 day -1 (n =
3; Table 3) to be a realistic estimate for sediments in the oxygen minimum zone. This SRR
is 50 to 300-fold higher than SRR measured on outer continental shelf sediments (150-
500 m water) outside upwelling areas (IVANOVet al., 1980; BATTERSBYand BROWN, 1982).
This difference is probably explained by the high flux of organic matter to the sediment due
to intense primary production in the upwelling waters. It is therefore not surprising that
sulfate reduction rates are as high as observed in estuaries and coastal marine sediments
(SKYRING, 1987; CANFIELD,1989).
To evaluate the importance of sulfate reduction for the carbon cycling in the Peruvian
upwelling system the sulfate reduction rates were compared to rates of water-column
primary production. Using an average SRR of 20 + 11 mmol SO42- m -2 day -1, and
assuming that 2 moles of CO2 are formed for each mole SO42- respired, the sulfate reducers
are able to remineralize (20-12-2) 480 + 260 mg C m -2 day -1. Primary production ranges
from 1 to 4 g C m -2 day -1 in most studies (LEE and CRONIN, 1982; HENRICHS and
FARRINGTON,1984), although RYrHERet al. (1971) have reported values up to 11.7 g C m -2
day -1, the highest ever reported for this area. The sulfate reducers thus remineralize
9-29% (0.48 + 0.26/2.5) of the total primary production of which 10-15%, based on
sedimentation trap measurements, was estimated to sink to the sea bed (LEE and CRONIN,
1982; HENRICHSand FARRINGTON,1984; ROWE,1985). However, this does not correspond
with the observation that about half the organic carbon is permanently buried in the
sediments (Figs 2-5). The sedimentation, though, may change from year to year. The
sedimentation trap measurements are also sparse and these measurements might not
correspond with the sedimentation on the stations investigated during this study. Organic-
rich sediments could also be transported downslope, as the bottom slope in this area is
about 1:10. A higher amount of organic carbon will be available in this way for
mineralization than can be calculated from the sedimentation of the local primary
production. A nonsteady-state deposition of this kind was proposed by HENRICHS and
FARRINGTON (1984) from 21°pb observations. The downslope transportation may also
explain the almost uniform SO42- gradients in the top 10-20 cm of the sediment in the
oxygen minimum zone (Figs 2-5). No bioturbation was observed on these stations and
diffusion alone cannot explain the SO 2- concentrations measured. It is, however, evident
from this study that sulfate-reducing bacteria must play a most important role in this
sediment ecosystem.

Acknowledgements--I thankJohnW. Farringtonfor the invitiationto participatein the PUBSI, 1987expedition

supported by Grant OCE-85-09859 U.S. NationalScienceFoundation. I also thank the officers,crew, and the
366 H, FOSSING

scientific party of R.V. Moana Wave for their contributions to the success of the sampling program. Thanks are
especially due to Eben Franks and Douglas C. Nelson for their help during subsampling. Else Bundgaard Frentz
carefully assisted with the analytical work and Bo Barker Jcrgensen reviewed the manuscript and provided
helpful suggestions. Thanks to two anonymous reviewers for comments on an earlier version of this manuscript.
Funding was provided from the Danish Natural Science Research Council, Grant nos 11-5736 and 81-5671.

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