AEROBIC DECOMPOSITION OF SEDIMENT AND

DETRITUS AS A FUNCTION OF PARTICLE SURFACE

AREA AND ORGANIC CONTENT
Bawy T. Hargrad
Freshwater Biological Laboratory, The University of Copenhagen, Hillergd, Denmark

ABSTRACT
Oxygen uptake by microbial populations on mud, sand, and various types of detritus was
measured in short-term experiments in aerated water at 20C. Sample size had no effect
on oxygen consumption per unit weight, but stirring increased uptake. A 2% Formalin
solution completely stopped biological uptake of oxygen by microorganisms on sand and
detritus; lake muds showed various degrees of chemical uptake of oxygen.
Microorganisms on dead Phrugmites leaves consumed oxygen at an increasing rate during
the first few days of decomposition, followed by a decline to a rate comparable to uptake
by freshly collected detritus. Limnaea feces initially consumed oxygen three times more
rapidly than detritus, but after 5 days the rates were equal.
Detritus consumed up to three orders of magnitude more oxygen per dry weight than
sand; uptake rates were inversely related to particle diameter. The logarithm of oxygen
uptake was directly related to the logarithm of particle organic content. Particulate oxygen
uptake in this and previous studies fell between 0.1 and 10 mg 02 (g organic matter)-l hr-l,
a rate inversely related to particle diameter. Oxygen uptake per unit weight by particles
of ashed mud and sand exposed for 24 hr to decomposing detritus in a nutrient solution
and transferred to fresh solution was inversely related to particle size and similar to rates
measured with freshly collected samples. On an areal basis all particles consumed between
0.01 and 1.0 x lo-’ mg O2 cm-l hr-I.
The negative linear correlation on logarithmic axes of surface area, organic carbon and
nitrogen and bacterial plate counts with sediment particle size is similar to that observed
for measures of oxygen uptake. Bacteria cover only a few percent of particle surfaces. This
may result in the narrow range of measures of microbial community respiration on an
areal basis.

INTRODUCTION oxygen uptake by detritus, mud, and

Aquatic sediments and detritus consume
oxygen when stirred in aerated water, The
oxygen uptake is an integrative measure
of all oxidative processes occurring in the
sample, both chemical and biological.
Reducing substances are usually rapidly
oxidized. Respiration of the organisms as-
sociated with particles is probably pri-
marily bacterial, although algae, protozoa,
and fungi may also contribute. Whatever
populations are involved, measurements of
oxygen uptake reflect the metabolism of
communities of microorganisms involved
in the decomposition of natural substrates.
No measure is provided, however, of an-
oxic microbial decomposition.
Many workers have measured rates of
l Present address: Fisheries Research Board of
Canada, Marinc Ecology Laboratory, Bedford In-
stitute of Oceanography, Dartmouth, Nova Scotia,
LIMNOLOGY AND OCEANOGRAPHY
sand sediments. Waksman and Hotchkiss
( 1938)) Teal ( 1962), and Rybak ( 1969)
suggested that oxygen uptake is related to
the organic matter available for decompo-
sition; Odum and de la Cruz (1967) and
Fenchcl (1970) d cmonstrated an invcrsc
relation between detritus particle size and
oxygen consumption. Chemical oxidation
of reduced subsurface sediment may also
be related to the concentration of reduced
substances (Gardner and Let 1965; Har-
grave 1972).
Teal (1962) and Odum and de la Cruz
( 1967) found approximately similar rates
of oxygen consumption by Spartina detri-
tus. Thalassia decays much faster than
does Spartina (Wood et al. 1969)) yet oxy-
gen uptake by detritus from these plants,
freshly collected or artificially prepared, is
similar ( Fenchel 1970). Lake Esrom scdi-
mcnts consume oxygen at rates similar to
583 JULY 1972, v. 17( 4)
584 BARRY
those observed with salt marsh mud (Har-
grave 1972).
I here compare rates of oxygen uptake
by sediment and detritus particles of dif-
ferent sizes and organic content, These
results, combined with previous measurc-
ments, show an inverse relation between
particle size and oxygen consumption. The
importance of surface area and organic
content is examined by calculating oxygen
uptake of natural and recolonized parti-
cles on a surface area basis. A possible
mechanism causing similarities and differ-
ences in rates of sediment oxygen con-
sump tion is suggested.
I gratefully acknowledge a NATO Sci-
ence Fellowship provided by The National
Research Council of Canada, Professors E.
Stecmann Nielsen and T. Fcnchel kindly
read the manuscript and offered com-
ments. Mr. S. Bordon calculated the re-
gression lines.
MATERIALS AND METI-IODS
Sediment and detritus from freshwater
sources were collected from eutrophic
Lake Esrom and Frederiksborg Castle
Lake, from the humic-acid brown-water
Store Grib Lake, all located in northern
Zealand (Denmark) (Berg 1938), and from
Grane Lang Lake, an oligotrophic lake in
Jutland (Whiteside 1970). Other samples
were taken from Hclsing@r, Kronborg, and
Julebaek beaches along the northern @re-
sund near Helsinger ( Fenchel 1969).
Either a Kajak corer (Brinkhurst et al.
1969) or a glass cylinder pushed by hand
into shallow sand sediment was used to
take undisturbed cores of mud and sand.
In the laboratory, samples of mud and fine
sand sediment wcrc taken from the surface
of the cores by pipette, mixed, and drained
on filter paper. About 0.1 g wet wt was
removed from the filter paper with a
spatula and added to 28-ml Pyrex BOD
bottles filled by siphon with filtered aer-
ated water from the collecting area. Large
pebbles, with a total weight of up to 5 g,
were added to bottles with forceps. Detri-
tus collected by hand from beach margins
T. I-IARGRAVE
was used immediately. Disks (l-cm diam)
were cut from larger pieces of decaying
vegetation with a cork borer and lo-20
mg (dry wt) were placed in bottles. In-
cubations lasted l-2 hr in the dark at 2OC,
and bottles were rotated (4 rpm) on their
long axis to prevent oxygen depletion
around sedimented particles. All experi-
ments were carried out during summer
and early fall when field temperatures
ranged between 10 and 22C.
A buffered (pH 6.8) solution of 2% For-
malin was used to fill some BOD bottles.
ZoBell and Brown (1944) found that a
0.25% Formalin solution stopped bacterial
respiration, so oxygen consumed by parti-
clcs in this solution was assumed to be due
to chemical uptake. Additional blank bot-
tles corrected for small amounts of oxygen
uptake by the Formalin.
Dissolved oxygen was measured by a
micro-Winkler technique (Fox and Wing-
field 1938) with samples taken in 5-ml
syringes; a solution of 1% NaN3 was added
to the NaI solution to prevent interference
from reduced substances. Syringe samples
of supernatant were taken after settling
of particles. Mud particles, in particular,
appear to bind the iodide liberated dur-
ing the Winkler reaction and thus reduce
the apparent oxygen concentration. For-
malin soIutions were titrated slowly over
10 min for accurate end-point determina-
tions. Measurements of dissolved oxygen
were accurate to at least 0.1 mg/liter.
Changes as smalI as 1% of total oxygen
present could be measured; changes were
usually much greater, but concentrations
were never reduced more than 30%. Cor-
rections were made for volume displace-
mcnt throughout.
Mud and detritus were filtered onto pre-
ashed and preweighcd glass-fiber filters,
dried ( 8OC for 24 hr ), and weighed. Sand
grains and pebbles were dried in pre-
weighed crucibles. Organic matter was
determined by loss on ignition at 550C for
3 hr; carbonate dissociation is avoided at
this temperature if ashing times are not
long (Rybak 1969). Particle size (mean
value of greatest dimension of randomly
 DECOMPOSITION OF SEDIMENT
chosen particles) of sand grains was deter-
mined by microscopic examination; at least
100 were examined when their diameter
was less than 500 p. All pebbles greater
than l-mm diameter in the bottles were
measured and the mean diameter dcter-
mined, Mud sediments consisted entirely
of silt and amorphous organic debris that,
with gentle agitation, passed through a
nylon screen with 100-p-mesh openings
but clogged screens of 20 p; an average
particle diameter of 50 p was assumed.
The detritus from decaying vegetation,
when not cut with a cork borer, was mea-
sured wet on graph paper or under a
binocular microscope.
Fecal pellets were collected from three
common benthic invertebrates in Lake Es-
rom. Adult Ilyoclrilus hummoniensis ( Oli-
gochaeta) and fourth instar Chironomus
anthracinus (Diptera) larvae were held in
freshly collected profundal lake mud, and
fecal pellets were collected from petri
dishes in which several animals had been
held overnight. Limnaea palustris ( Mol-
lusca ) , collected with decaying Phragmi-
tes communis, browses epiphytic material
and produces abundant feces. All fecal
pellets were kept in gently aerated lake
water when not used within a few hours,
Rates of oxygen consumption per unit
weight were compared on a surface area
basis by dividing rates of oxygen uptake
by the number of square centimeters of
sediment. Area was calculated for detritus
particles larger than 1 mm before drying
by doubling that traced on graph paper.
Values for mud and detritus with a par-
ticle diameter less than 1 mm were taken
from Fenchel’s (1970) estimate of total
surface area per gram dry weight of Tha-
lassia detritus. Sand grain and pebble sur-
face area was estimated from a relation
between mean sand grain size and “in-
ternal surface” arca derived by Fenchcl
( 1969), based on 10 cm3 of sample and
the assumption that sand grains were
spherical. If the specific gravity of sand
(quartz) is assumed to be 1.6 (Kaye and
Laby 1956), then the surface area per
gram dry weight for a given sand grain
AND DETRITUS 585
diameter is Fenchel’s internal surface area
divided by 16.
Recolonization expcrimcnts were on
mud, sand, and pebbles from Lake Esrom
sterilized by ashing at 550C for 3 hr. A
small piece of decomposing Phragmites
leaf in a solution of 1% each of glucose,
peptone, and K&P04 and mud-water ex-
tract (1 kg of wet profundal mud auto-
claved with 1 liter of Lake Esrom water)
was used for inoculum, with controls of
sterilized particles in autoclavcd lake wa-
tcr without detritus. After 24 hr of aera-
tion, control and inoculated particles were
rinsed and placed in freshly filtered water,
and oxygen uptake was measured.
RESULTS
Experimental variables
Lake Esrom mud between 1 and 20 mg
( dry wt ) per bottle gave no significantly
different rates of oxygen uptake per gram
dry weight. Eight samples of surface sedi-
ment taken from four profundal mud cores
during June consumed 0.22 * 0.04 mg 02
g-l hr-l regardless of sample size. Samples
of sand grains (250-p diam), from a beach
surface in Lake Esrom, of dry weights
up to 1 g showed similar reproducibility
(220%). Two replicate samples were then
used for each determination and the mean
value recorded. Short incubations, l-2 hr,
ensured sufficient oxygen yet gave mea-
surable changes in concentration. The
rapid oxygen uptake by some sediment
and detritus particles necessitated agita-
tion of samples. Profundal mud from Lake
Esrom consumed 0.32 mg 02 g-l hr.-l when
bottles were slowly rotated and 0.08 when
standing. Oxygen uptake of beach sand
decreased by 15% when it was not stirred.
Relative importance of chemical
and biological oxidation
Oxygen uptake by sand and detritus
from beaches was always completely
stopped by 2% Formalin; oxygen uptake
by subsurface sediment from below 2 cm
in cores from Store Grib Lake and Lake
Esrom was unaffected by Formalin treat-
586 BARRY T. HARGRAVE

TABLE 1. Effect of Formalin treatment (+ = with, - = without) on oxygen uptake by surface (up-
per 5 mm) mud from four Danish lakes at various times of the year. All values are means of at least
two separate determinations

Oxygen uptake
- + by chemical Biological Sediment mg 0,
oxidation respiration org matter (g org
(mg 0, g-1 hr-1) (%I (mg 0, g-1 hr-1) (%I matter)-1 hr-1

Esrom
15 Jun 0.75 0.44 59 0.31 39.6 0.78
13 Nov 1.48 0.87 59 0.61 40.0 1.52
15 Dee 1.14 0.35 31 0.79 39.9 1.98
x 1.43
Store Grib
5 Sep 0.74 0.48 65 0.26 72.0 0.36
Freclcriksborg Castle
22 Sep 1.34 0.73 54 0.61 45.0 1.36
Grant Lang
15 Mar 0.10 0.01 10 0.09 21.3 0.43

ment, indicating that oxidative processes
were chemical, Surface muds (upper 5 mm
as taken by pipette) from various lakes,
however, showed varying degrees of sensi-
tivity to poisoning (Table 1). Oxygen was
present ( 3-5 mg/liter ) over all sampled
cores. Data from Lake Esrom show con-
siderable seasonal variation in total oxygen
uptake and the amount susceptible to For-
malin poisoning.
Particle size and organic content
Oxygen uptake by sand grains, detritus
particles, and mud sediments was differ-
ent per unit weight and each varied on the
basis of mean particle diameter (Fig. 1).
With the logarithmic transformation of
both axes, Fig. 1 reveals a linear, inverse
relation between particle diameter and ox-
ygen uptake. Values for mud were omitted
from the regression calculations because of
the inaccurate estimation of particle size.
Oxygen uptake for a given particle diam-
eter was two to three orders of magnitude
higher for detritus than for sand. Sand
grains and pebbles contained less than 1%
organic matter while mud and detritus
particles contained 40-90% organic matter.
After logarithmic transformations, oxygen
uptake was linearly related to organic con-
tent (Fig. 2).
The effect of differences in percentage
of organic matter in various types of scdi-
ment was standardized by expressing oxy-
gen uptake on an ash-free weight basis.
In sand and detritus this measure reflects
the metabolic activity of communities of
microorganisms since these substrates
showed no chemical oxidation. Mud par-
ticles did undergo chemical oxygen uptake
and this has been subtracted in the calcu-
lations of biological respiration in Table
1. The range of values of oxygen uptake
by various particles was reduced by these
calculations, and the effect of particle size
was enhanced (Fig. 3). Oxygen uptake by
all material fell within the range [O.l-10.0
mg 02 (g org matter)-l hr-l] observed for
detritus ( Fig. 1) largely because values
for oxygen uptake by sand grain commu-
nities, when expressed on an ash-free ba-
sis, were raised. Although differences are
large, organic matter in various muds was
oxidized at much lower rates than detritus
particles of similar size.
Stage of decomposition
Fresh snail fecal pellets (about 0.5-mm
diam and 2 mm long) contained 80-90%
organic matter. This always increased as
they aged, probably from organic matter
removed from solution by microorganisms
colonizing their surfaces, although it could
also result from leaching of inorganic salts.
 DECOhXTi’OSITION OF SEDIMENT AND DETRITUS 587

bacteria

,
,
/
/
/
/
/
/
sand ,
,

0
0
0
. x
-4 -I 0 8c,““I 1 8 I II”11 1 18““‘I ’ “A?
0 1 2 3 4
loglo particle diameter
(I-r)
FIG. 1. Relation between mean particle diam-
eter and oxygen uptake per gram dry weight of FIG. 2. Relation between percent organic mat-
sand, mud, and detritus from various aquatic ter and oxygen uptake per gram dry weight of
habitats. different particles described in Fig. 1. Regression
Solid circle-detritus from Lake Esrom shore line, calculated omitting bacteria and mud sam-
(1, mean value for five species of deciduous tree ples, y = -2.15 + 0.90x (r = 0.91, sE of slope
leaves; 2, unidentified reed detritus; 3, Phrugmi- 0.075, significant at P = 0.05).
tes leaves); solid triangle-invertebrate feces (l-
day-old ) ( 1, Limnaea palustris; 2, Ilyodrilus
hammoniensis. 3, Chironomus anthrucinus); solid days the rate of oxygen uptake declined
square-sand and unknown seaweed detritus from
Julebaek beach; cross-sand, detritus, and mud
to 0.55 mg O2 g-1 hr-l, followed by a
from Store Grib Lake; open triangle-Frederiks- slow decrease to 0.25 after holding for 20
borg Castle lake mud; point-Grane Lang Lake days. The decline in oxygen uptake was
profundal mud; circled cross-IIelsinggr beach accompanied by successions of communi-
sand; circled point-Kronborg beach pebbles;
open square-sediment bacteria (Pseudomonas
ties of microorganisms associated with the
spp.) cultured from Lake Esrom surface profundal fecal material. Ciliates, abundant after the
mud; open circles-Fenchel (1970), Thalassia de- first day, declined after the fifth. Fungal
tritus; squared points-Odum and de la Cruz hyphac grew rapidly and changed the
( 1967), Spartina detritus. pellets from brown to white within 12 hr.
Regression lines-detritus: y = 1.45 - 0.52 x ( r
= 0.65, SE of slope 0.17); sand: y = -0.20 - 0.85 Phragmites leaves, collected dry from
x (r= 0.95, SE of slope 0.07). Slopes are signifi- dead plants in November, were placed in
cantly different from zero (P = 0.05). Data for aerated lake water in the laboratory, Disks
mud, bacteria, Thalassia detritus ( Fenchel 1970))
and Spartina detritus (Odum and de la Cruz 1967)
of leaf material tested immediately con-
not included in regression calculations. tained 82.5% organic matter and consumed
0.22 mg 02 g-l hr-l (Fig. 4). After 3 days
Oxygen uptake by microorganisms on fe- oxygen uptake had increased to 1.0 mg
cal pellets used within 6 hr of production 02 g-l hr-l and leaf material contained
was three times faster than when pellets 90% organic matter. During the next 2
were first held for 12 hr (Fig, 4). After 3 days there was little change in organic
588 BARRY T. HARGRAVE
log ,O particle diameter
0-l)
FIG. 3. Relation between mean particle diam-
eter and oxygen uptake per gram organic matter
(calculated from Figs. 1 and 2 ). Symbols as in
Fig. 1. Regression line, calculated omitting mud,
bacteria, and data (line joining points) from Odum
and de la Cruz (1967), y = 2.07 -0.69x (r = 0.79,
SE of slope 0.10, significant at P = 0.05).
content ( 90.5% ) , but oxygen consumption
fell to the original level. Oxygen uptake
remained constant over the following 34
days, while organic matter slowly in-
creased to 98.5%.
In a similar experiment with Phragmites
leaves collected as wet detritus from be-
side Lake Esrom, oxygen uptake, initially
0.5 mg 02 g-l hr-l, decreased steadily,
while organic matter increased from 94.3-
98.0%. The final constant rate of oxygen
consumption by this material was similar
to those by freshly wetted leaves and
Limnaea feces at the end of the holding
period (Fig. 4). This value (0.25 mg 02
g-1 hrl) falls within the range observed
for detrital material from other sources
(Fig. 1).
c 0 fecal pellets
“,~~~~~‘, Phragmites detritus
1.0
0 10 20 30 40
Time (days)
FIG. 4. Changes in oxygen uptake by Limnuea
feces and Phragmites detritus held at 20C in aer-
ated Lake Esrom water. Samples removed for use
in short-term experiments at various times. a-
Detritus prepared from plant leaves collected dry;
b-detritus wet and partly decomposed when
collected.
Previous measurements
Odum and de la Cruz (1967) and
Fenchel ( 1970) related oxygen uptake to
particle size of Spartina and ThaZussiu de-
tritus. Their results, plotted in Fig, 1, are
of the same magnitude as mine for detri-
tus from different sources.
I have used organic contents of sedi-
ment and detrital material reported by
various workers as a basis for comparing
rates of oxygen uptake per unit dry weight
(Fig. 5); sewage particles, artificial stream
periphyton, aquatic vascular plants, and
aquatic bacteria are included for compari-
son. When expressed on logarithmic axes,
these data show the same trend and span
the same range as those from this study
(Fig. 2). Odum and de la Cruz ( 1967)
measured both organic content and parti-
cle size of decomposing Spartina detritus.
 DECOMXPOSITION OF SEDIUENT AND DETRITUS 589

m mg O2 g-’ hr-’
0 percent organic matter
m mg O2 (g organic matter )-’ hr-’
m mg o2 x10-~ cm-* hr-’

particle diameter CJI)

mud sand pebbles

FIG. 6. Mean oxygen consumption and organic

log,o percent organic content of triplicate samples of ashed mud, sand,
matter and pebbles from Lake Esrom after 24 hr of
FIG. 5. Comparison of organic content and ox- exposure to Phragmites detritus in a nutrient-cn-
ygen uptake per gram dry weight for different riched solution. Control samples, not exposed to
types of sediment and detritus, aquatic vascular detritus and nutrients, contained no mensurable
plants, periphyton communities, and bacterial cells organic matter and consumed no oxygen.
from previous studies. Circles indicate average
values for measurements at about 20C. Lines rep-
resent range of published observations. Oxygen uptake, on an ash-free basis, fol-
l-Johnson ( 1936 ) , resting marine bacteria; lows the trend of my data and falls within
2-ZoBell and Stadler (1940), multiplying lake the same order of magnitude when com-
bacteria; 3-Liagino and Kusnetzov (cited in Zo-
Bell and Stadler 1940), lake bacteria; 4-Har-
pared with particle size ( Fig. 3).
grave (1969), estimated value for lake sediment
bacteria; 5-Gessner ( 1959), various aquatic vas- Particle surface area
cular plants; 6-McIntire ( 1966)) periphyton All particles considered in this study
communities in artificial streams; 7-Teal (1962),
Spartina detritus during 4-week decay period; 8- consumed between 0.01 and 1.0 x 1O-3 mg
Odum and de la Cruz ( 1967), Spartina detritus; 02 cm-2 hr-l with the exception of freshly
9-Fenchel ( 1970), Thalassia detritus. produced Limnaea feces (Table 2); oxygen
N-Water Pollution Research Rep. ( 1968),
consumption of fcccs fell to within the
suspended solids in treated sewage effluent; ll-
DiSalvo ( 1971), coral reef regenerative sediment; range observed for all other particles
12-I&rnerblad (1930), mean value for 14 Swed- within 3 days. Dctri tus particles con-
ish lakes ( assumed 2 g dry wt/flask); 13- sumed more oxygen on an arcal basis than
Miyadi (cited in ZoBell 1946), lake sediments;
14-Waksman and Hotchkiss (1938), marine sedi- pebbles, sand, or mud. There was no con-
ments from Woods Hole; 15-Kato ( 1956), ma-
rine bottom sediments from the northern Japan
Sea; 16-Teal and Kanwisher (1961), surface salt-
marsh mud; 17-Gardner and Let ( 1965), Lake (1969), black clots, Polish lakes of varying trophic
Mendota sediment (initial 3-day rate); 18-Rybak type.
590 BARRY T. IIARGRAVE

TADLE 2. Comparison of mean particle diameter recolonized pebbles, however, was an or-
(p), estimated surface area per gram dry weight
der of magnitude higher than that of
(A), and oxygen uptake per unit surface area (C)
for various types of particles. Data fur oxygen freshly collected material. There were few
uptake per gram (B) from Fig. 1 protozoa in the detritus used for inocula-
tion and thus oxygen consumption prob-
Particle
and source A B C
ably reflects only bacterial respiration.
P
No organic matter remained on particles
Pebbles after ashing, but after 24 hr of exposure
Kronborg beach 9,000 1.2 0.00012 0.10 to the nutrient solution and detritus par-
7,800 1.5 0.00030 0.20
ticles a loss on ignition was measured in
5,340 1.9 0.00655 0.2.9
4,000 2.7 O.OC@70 0.26 all samples (Fig. 6). The percentage or-
Esrom beach 1,914 5.6 0.0012 0.21 ganic matter was inversely related to par-
1,260 6.5 0.0011 0.17 ticle size but all particles contained an
Sand order of magnitude less organic matter
Esrom beach 550 21.9 0.0058 0.26 than naturally occurring samples of simi-
3G6 31.2 0.0028 0.09 lar type.
Helsing@r beach 350 32.0 0.0046 0.14 Oxygen uptake per gram organic matter,
185 62.,5 0.0064 0.10 which ranged from 9-28, was not related
Julebmk beach 248 50.0 0.0086 0.17
to particle diameter (Fig, 6). Oxygen
Lake mud consumption per unit surface area in-
Esrom 50 6,000 0.86 0.14 creased with increasing particle size; the
Store Crib 50 6,000 0.74 0.12 range of values was within that observed
Grane Lang 50 6,000 0.10 0.02
Frederiksborg Castle 50 6,000 1.34 0.22
with natural particles (Table 2).
Detritus DISCUSSION
Esrom shore debris - - 0.33 0.30
Wet elm leaves - - 0.18 0.42
The enumeration of microorganisms can-
Phragmites - - 0.24 0.42 not serve as a measure of their importance
E quisetum - - 0.10 0.26 in proccsscs of decomposition. There may
Limnaea feces, fresh 5001 650 3.50 5.38 be no clcarcut relation between numbers
3-day-old 500 650 0.55 0.85 and metabolic rate. The dynamic nature
Spartina 239 1,050 0.63 0.60
(Odum and de 150 1,800 1.62 0.90 of microbial populations also makes esti-
la Cruz 1967 ) 64 4,200 1.81 0.43 mates of population size of limited use;
Thalassia 1,000 420 0.40 0.95 grazing may stimulate or retard cell divi-
(Fcnchel 1970) 400 850 0.60 0.71 sion and substrate changes determined by
100 2,500 1.05 0.42
species succession may continuously affect
population size. These limitations prob-
ably account for the lack of correlation
sistent relation between oxygen uptake per between sediment organic content and
square centimeter and particle size. indices ( plate counts) of bacterial biomass
(Cooper et al. 1953; Volkmann and Oppen-
Recolonization experiments heimer 1962; Anthony and Hayes 1964).
Previously ashed pebbles, sand, and Also, the amount of total organic matter
mud, held in sterilized water for 24 hr, in sediment does not indicate its availa-
showed no measurable oxygen consump- bility to cithcr microorganisms ( Waksman
tion. After 24 hr of exposure to Phragrni- and Hotchkiss 1938) or invcrtcbratcs (Har-
tes detritus in an enriched medium, rinsed grave 1970a).
samples of mud and sand consumed oxy- Measurement of oxygen consumption
gen at rates inversely related to particle appears to be a more useful basis for com-
size and in the range of similarly sized paring microbial activity in different types
particles with natural communities of mi- of sediment and detrital material. Hayes
croorganisms (Fig. 6). Oxygen uptake by and MacAulay (1959) suggest that oxygen
 DECOMPOSITION OF
uptake by mixed mud may represent an
integrative measure of lake productivity.
Although they measured oxygen consumed
by restratified mud-water systems, there
was a linear relation between oxygen up-
take and “lake quality index,” a value
derived from an estimate of fish yield.
Experimental variables
The interpretation of oxygen uptake
necessitates standardization of those exper-
imcntal variables which might affect mea-
sured rates. For example, Waksman and
Hotchkiss (1938) found shaking to have
no effect, but total oxygen uptake de-
creased with the amount of sediment per
bottle; I observed opposite effects of shak-
ing and sample size. They left sediment
standing for up to 30 days, stirring it by
hand several times daily. In long-term
experiments, oxygen dcplction around set-
tled particles would lower the rate of oxy-
gen uptake. The effect would be greater
with larger samples ( Kato 1956). Oxygen
uptake by mud sediments, because of the
small particle size and often high propor-
tion of chemical oxidation ( Table I), may
bc more affected by agitation and size of
sample than sand sediments. The small
range of weights used hcrc (1-28 mg dry
wt/28 ml) may not have been sufficient
to show such an effect.
Sample variability
Changes in rates of oxygen consumption
by sediment and detritus with time can
be expected as easily oxidized substances
disappear. Previous studies with various
types of scdimcnt (Anderson 1939; Kato
1956; Teal and Kanwisher 1961; Gardner
and Lee 1965; Hamilton and Greenfield
1967; Rybak 1969; Fenchel 1969; DiSalvo
1971) all show a time-dependent decrease
in oxygen consumption although the time
scale is different. The immcdiatc effect
may represent oxygen uptake by reduced
minerals, such as ferrous iron and ferrous
carbonate, and microbial decomposition of
easily oxidized organic substances. Long-
term slow rates of oxygen consumption
may bc due to acid-insoluble iron sulfide
SEDIMENT AND DETRITUS 591
minerals (Gardner and Lee 1965) and
slow microbial oxidation of organic matc-
rial such as lignin and humic complexes.
Whatever the cause of the time-dependent
decrease in oxygen uptake, short-term ex-
pcrimcnts give maximum rates of oxygen
consumption, which might reflect rates oc-
curring in nature if samples were taken
from well-aerated areas.
The depth from which a sample is taken
can be important. The variable proportion
of chemical oxidation in lake muds (Lon-
ncrblad 1930 and Table 1) differs from
marinc sand sediment where sterilization
completely stopped oxygen uptake (Waks-
man and Hotchkiss 1938). The absence
of chemical oxidation of beach sand and
detritus probably results from the well-
oxygenated conditions on wave-washed
beaches where few reducing substances
exist ( Fenchel 1969). Usually, however,
sediments contain an oxidized surface
layer overlying anoxic reduced sediment
( Hayes 1964; Fcnchcl and Ricdl 1970).
Oxygen uptake by subsurface sediment
may bc cntircly chemical ( Moore 1931))
but thin layers of surface sediment show
varying proportions of chemical and bio-
logical oxygen uptake (IIargrave 1972 and
Table 1) . Biological oxidation accounted
for an average of 30% of oxygen consumed
by salt marsh mud (Teal and Kanwisher
1961), with no clear effect of depth,
Samples of sediment and detritus held
in acrated water rapidly become dcoxy-
genatcd and anaerobic bacteria appear.
This is not surprising since oxygen uptake
by aquatic bacteria at 20C may range
from NO-500 ~1 02 mg-l hr-l ( Fig. 5). A
1-g sample of detritus at 20C consumes
about 1 mg 02 hr-l; thus, all of the oxygen
in 100 ml of water could bc consumed
within 1 hr. The rapid rate of oxygen con-
sumption by bacteria in sediment may de-
oxygenate bottom deposits ( ZoBell 1946).
Dissolved oxygen may only penetrate the
top few millimeters of mud sediments not
exposed to water turbulence ( Hargrave
1972). Oxidative decomposition in such
habitats must bc limited by oxygen supply.
The supply of easily oxidized organic
592 BARRY T,
matter must also affect rates of sediment
oxygen uptake. The higher rates of oxy-
gen uptake by surface sediment in Lake
Esrom in November and Dcccmber may
reflect the decomposition of organic mat-
ter newly deposited after phytoplankton
blooms ( Table 1).
The stage of decomposition of detritus
is important. Organic content of fcccs may
increase or decrease with time (Newell
1965; Hargrave 1970b ) , indicating bactc-
rial colonization ( Fenchel 1970). Feces
were rapidly colonized (Fig. 4) and within
4 days, oxygen uptake decreased from 3.5-
0.4 mg 02 g-l hr-l. Phragmites detritus
did not reach as high rates of oxygen up-
take as feces but also showed a decrease
with time. Presumably initially high rates
result from an increase in microbial respi-
ration as labile substrates, such as mucous
secretions are utilized. Grazing organisms
such as protozoa, which appeared in both
feces and detritus during initial maximum
rates of oxygen uptake, would also con-
tribute to increased oxygen consumption.
Teal (1962) observed a peak in oxygen
uptake in freshly produced Spartina dctri-
tus during the first week of decomposi-
tion, similar to that noted with Phragmites
(Fig. 4). Explanations for differences in
the maximum rate obtained by Spartina
detritus ( 14 mg O2 g-l hr-l ) , Limnaea fe-
ccs (3.5), and Phragmites detritus ( 1.0)
during the initial period of decomposition
as well as the varied final rates of oxygen
uptake (0.25 mg O2 g-l hr-l for feces and
Phragmites, 2.0 for Spartina) may reflect
the species of microorganisms involved
and the residual substrates.
The difficulty of following decomposi-
tion by only observing changes in organic
matter is illustrated by the increase in
organic content of Phragmites during de-
composition, in contrast to Spartina (Teal
1962 ) . Changes in organic content in-
cludc changes in both organic substrate
and communities of colonizing microorga-
nisms, but changes in rates of oxygen con-
sumption arc more easily interpreted as
they reflect the metabolic activity of all
organisms associated with aerobic de-
HARGRAVE
composition. The time-course of detritus
decomposition as measured by oxygen up-
take seems similar for different substrates.
It is not clear why different particles of
a similar size should have even an approx-
imately similar rate of oxygen uptake
(Fig. 3). Decomposition processes in dc-
tritus and mud, where the substrate is
metabolized, may differ from those on
sand and pebbles where mineral surfaces
serve largely as a source of attachment.
If, however, only organic matter adsorbed
to surfaces is available to heterotrophic
bacteria ( ZoBell 1946)) oxygen consump-
tion by microorganisms associated with
sand and pcbblcs also reflects substrate
decomposition.
Oxygen uptake by mud particles tends
to be lower than might be expected on
the basis of its particle size relative to that
of detritus, and at least some of the oxy-
gcn is used for chemical oxidation. The
somewhat arbitrary estimate of mud par-
ticle diameter makes comparison on the
basis of particle size difficult. Organic
matter in lake sediment ranged from 20-
70%; detritus always contained more than
70%. The substrates in mud may bc less
easily oxidized. Grane Lang Lake mud
consumed the least oxygen ( 0.10 mg 02
g-l hr-l ) and also had the lowest organic
content (21.3%) of the lake sediments ex-
amined. Low annual primary production
in this lake means that its surface scdi-
ment is older than that of the more pro-
ductive Lake Esrom or Frcderiksborg
Castle Lake (Whiteside 1970). Rybak
(1969) reported that lake sediments that
consumed less oxygen are poorer in or-
ganic matter; data summarized from other
work (see Fig. 5) shows a similar trend.
These observations support Hayes and
MacAulay’s ( 1959) suggestion for sedi-
mcnt oxygen uptake as an index of lake
productivity.
Particles of various types have diffcr-
ent rates of oxygen uptake per unit sur-
face area, but the range of values is not
large (Table 2). Estimates of surface area
per gram for various particles span four
orders of magnitude, although oxygen up-
 DECOMPOSITION OF
take per square centimeter only has a
range of two. Surface area was only mea-
sured for large pieces of detritus, and the
values assumed for other particles from
previous studies could introduce errors;
however, the cstimatcs are probably accu-
rate enough for present calculations and
suggest some constancy in total commu-
nity metabolism per unit surface for very
different types of particles.
Recolonization experiments
Oxygen consumption per unit weight
and the adsorption of organic matter were
both inversely related to particle size in
recolonization experiments, but the dif-
ferences were not as great as would be
expected on the basis of surface area. Ox-
ygen uptake per unit weight and surface
area was within the range found with
freshly collected samples. All recolonized
particles had an approximaetly similar rate
of oxygen uptake on the basis of organic
content, higher than that found with the
smallest detritus particles used by Odum
and de la C*luz ( 1967). Sincc respiration
of rinsed particles was measured after
transfer to fresh nonenriched solutions, or-
ganic matter on the surfaces must have
undergone relatively rapid oxidation simi-
lar for each particle type; the organic mat-
ter and microbial populations adsorbed to
the various particles in these experiments
may also have been similar.
Recolonized particles should all have
similar rates of respiration per unit sur-
fact if oxygen uptake by communities of
microorganisms on particle surfaces is de-
pendent only on surface area, However,
uptake increased with increasing particle
size ( Fig. 6). There was a disproportion-
atcly high organic content on pcbblcs;
mud contained less organic matter than
expcctcd on the basis of its surface arca.
These differences may result from altcra-
tion of the surface characteristics of the
particles in ashing, Badcr (1962) did not
find simple adsorption phenomena bc-
twecn dissolved organic compounds and
mineral particles. Various sediments differ
in adsorbing properties for different spc-
SEDIMENT AND DETRITUS 593
ties of bacteria (Rubcntshik et al. 1936).
ZoBell (1943) noted a logarithmic increase
extending over 18 days of bacteria at-
tached to glass surfaces; 24 hr is probably
insufficient for populations of colonizing
microorganisms to reach a stable level of
density and metabolism.
Bacterial density on particle surfaces
Fenchel (1970) obscrvcd that both oxy-
gen uptake by Thalassia detritus and the
number of bacteria, small zooflagellatcs,
and diatoms wcrc proportional to surface
arca. The number of organisms per sur-
face area, about 3 x lo6 bacterial cells
cm-2 by direct counts, was always similar
on intact dead leaves, various sized parti-
cles, and artificial detritus after 4-6 days.
Tsernoglou and Anthony (1971) calculated
the density of bacteria by direct micros-
copy on freshwater sediment particles and
summarized previous estimates of bactc-
rial density on sand, pebbles, and soil
surfaces; depending on the source of the
sediment, there was one bacterial cell for
every 70-300 p2. If an average bacterial
ccl1 is assumed to cover 1 p2, then <I%
of the particle surface arca was colonized.
The bacterial density observed on detritus
by Fcnchel (1970) would cover about 3%
of the surface area.
Batoosingh and Anthony ( 1971) also
calculated bacterial density on pebbles di-
rcctly by fluorescent microscopy. Single
cells accounted for 70% of the bacterial
populations, When colonies were present
many seemed to be chance aggregations
rather than colonies, as the individuals
were generally quite separate from each
other. An analysis of the dispersion of
the direct counts showed that the bacteria
were slightly over-dispersed on the pcbblc
surfaces. There was an average of 1 cell
/300 /.L!
These calculations show, however ap-
proximately, that bacterial cells do not
completely cover these sediment and de-
trital particles. Other habitats, such as the
surface film of stagnant water, may have
higher ccl1 densities ( Fenchel, personal
communication), but natural sediment
594 BARRY T. HARGRAVE

tcrial numbers (Batoosingh 1964). Detritus,

more flocculant and with a higher organic
content than sand or pebbles, should of-
3
fer a more suitable substrate. Still, from
Fenchel’s ( 1970) observations, bacterial
2 cover in detritus is far from continuous.
Burkholder (1963) has shown that growth-
regulating substances are released by scdi-
mcnt bacteria. ZoBell (1943) described a
faintly staining film which surrounds bac-
terial cells attached to glass slides, two
or three times the dimension of the cells
themselves, whose size increases with age.
The nature of this material is unknown.
\\ Whatever the cause of spacing between
percent ‘. \
z
organic ‘* bacterial cells, only a fraction of the par-
-2- \ ‘. \
nitrogen ‘. ‘. ticlc surface arca is covered and the eco-
‘. logically available space is considerably
‘*.
-3 - mg O2 g-’ hr-‘x less than the total arca. Such a relation-
sand
ship bctwecn density and surface area
\ would impart some degree of similarity
-4 -I I I , 1\ to measures of community respiration per
0 1 2 3 4
unit area (Table 2). An upper limit to
loglo particle diameter
(J-0
cell density may be determined by avail-
able space, with metabolism per cell
FIG. 7. Comparison on logarithmic axes of related to the prcscncc of oxidizable sub-
mean particle diameter and internal surface area
strates, the two factors interacting to pro-
of sand (from Fenchel 1969), percent organic
carbon and nitrogen in beach sediments (from duce a relatively constant areal rate of
Longbottom 1970), bacterial plate counts in dif- community metabolism. This would ex-
ferent sediments (from ZoBell 1946), and mea- plain why although detritus consumes more
sures 0I particle oxygen consumption taken from oxygen per unit surface than do sand,
Figs. 1 and 3. Solid lines are calculated regres-
sion lines presented in the respective references.
pcbblc, or mud particles, all rates vary
The regression for bacterial numbers on particle within narrow limits.
diameter is y = 4.00 - 0.94x, T = 0.95. Dotted Longbottom ( 1970) reported that in
line indicates a slope of -1.0. sediments from gently sloping, moderately
well-drained beaches, both organic nitro-
particles of various types do not appear gcn and carbon content varied inversely
to bc heavily colonized. Cell counts per with median particle diameter after loga-
unit weight give a false impression of high rithmic transformations. The slope coeffi-
numbers; the spatial distribution of micro- cicnts, about -1.0, and the intercept values
organisms is given pcrspectivc when den- of the regression lines showed little sca-
sity is related to surface area. sonal change and were similar for sedi-
Many factors are involved in the spacing ments from diffcrcnt arcas. Fcnchel (1970)
of bacterial cells. Grazing and physical has also observed an inverse linear rela-
abrasion can remove cells. Algal and bac- tion (h = -1.0) bctwecn median particle
terial cells are usually found in depressions diameter and surface area of sand grains
and crevices of sand grains and pebbles on a logarithmic basis. These curves are
(Meadows and Anderson 1966), suggest- redrawn in Fig. 7. Apparently organic
ing physical removal from raised surfaces. matter adsorbed or attached as micro-
Particles removed from these influences, organisms to these sediment particles is
however, do not show an increase in bac- proportional to surface arca and the rcla-
 DECOMPOSITION OF SEDIMENT AND DETRITUS 595

tionship is approximately similar for sam- mining rates of oxidation of sediment and
ples from different arcas. detrital organic matter.
The relationship between sand grain The narrow range in rates of oxygen
size and oxygen uptake per unit weight uptake per square centimeter by different
also approaches a slope value of -1.0 (Fig. particles of a similar size suggests that
7)) perhaps reflecting the close correlation communities of microorganisms colonizing
between sand grain diameter and surface various surfaces may have similar rates of
area; mud and detritus particles are not metabolism. In aerobic habitats, coloniza-
spherical, so they need show no similar tion and succession may occur to the
relation. point where all of the oxygen supply is
Detritus oxygen consumption per unit fully used, Measurement of oxygen up-
weight is also inversely related to particle take by communities of microorganisms
size but with a slope less than -1.0 (Fig. associated with sediment and detritus par-
7). The relationship bctwccn particle di- ticlcs offers a simple method for investi-
ameter and external surface area per gram gating these processes.
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