Hydrobiologia 373/374: 203–216, 1998.

J.-C. Amiard, B. Le Rouzic, B. Berthet & G. Bertru (eds),

Oceans, Rivers and Lakes: Energy and Substance Transfers at Interfaces.
203
© 1998 Kluwer Academic Publishers. Printed in Belgium.

Phosphate mobility at the sediment–water interface of a Mediterranean

lagoon (etang du Méjean), seasonal phosphate variation

E. Gomez1 , M. Fillit2, M. C. Ximenes2 & B. Picot1

1 Département Sciences de l’Environnement et Santé Publique, UMR CNRS 5556, Ecosystèmes lagunaires, Faculté
de Pharmacie. 15 Av. C. Flahault, 34060 Montpellier, France
2 CEMAGREF, BP 5095, Montpellier, France

Key words: Phosphate, sediments, coastal lagoon, seasonal variations, P fractions

Abstract
The Méjean Lagoon (France) is a confined, shallow system, 0.7 m in depth, with a surface area of 747 ha. Its
sediments have a fine granulometry and are evenly distributed. The bathymetry and hydrodynamic behaviour of
the basin create two zones with restricted exchanges between them.
The western part (60% of the total surface area) is rich in dissolved phosphate (> 1 mg l−1 in summer) and
receives the majority of the phosphate (P) inputs from the watershed. The heavy macroalgal population consists of
Gracilaria throughout the year and Ulva in summer and autumn. Phytoplankton levels sometimes reach 100 µg
l−1 of chlorophyll a.
The sediment-water phosphate exchange varies with environmental conditions and macroalgal growth. P
mobility was studied on the basis of these characteristics.
Algal biomass and water characteristics were measured at 15-day intervals throughout 1993. A seasonal study
was conducted in 1994 to investigate which P fractions in the sediment were involved in the exchanges. The spring-
time rise in temperature reactivated decomposition of the macroalgal biomass that had accumulated in autumn. The
redox potential (EH ) fell as a result of this biological activity, leading to a decrease in the inorganic P fractions.
This P release accounts for the higher P water concentrations observed in the lagoon in summer.
The mobility of P fractions shows that the P stored in the sediments plays an active role in the dynamics of the
overlying water. Seasonal variations in these fractions explain the patterns of P storage and mobilization.

Introduction quantities of algae can produce anoxic conditions in

As human activity in the watersheds increases, an in-
creasing amount of waste from the continental land
mass enters the sea. Coastal lagoons act as an inter-
mediate recipient, protecting the marine environment.
They act as a sink for the pollutants. When large
amounts of nutrients (N and P) enter these stretches
of water, repeated algal blooms are observed and the
lagoons become eutrophic.
Coastal lagoons occupy a significant part (13%)
of the world’s shorelines (Nixon, 1982). These shal-
low ecosystems are usually very productive and play
an important role in the economic development of
the coastal zones. The decomposition of the large
the water column. The oxygen depletion results in
widespread mortality in the fish and farmed shellfish
populations. Massive algal growth has been observed
in various different eutrophic bays and lagoons: Saint
Brieuc Bay, the Po River Delta, the Venice Lagoon,
the Salses-Leucate Lagoon, the Prévost Lagoon, etc.
(Lieutaud et al., 1992; Pirou & Menesguen, 1992;
Viaroli et al, 1991; Sfriso et al., 1987; Boutière et al.,
1982).
To understand the internal functioning of these
ecosystems, two sites where nutrients are preserved
can be taken into consideration: the sediment and the
algae (macrophytes and phytoplankton).

Article: hy-ja28 Pips nr. 165699 (hydrkap:bio2fam) v.1.1

hy-ja28.tex; 2/09/1998; 19:22; p.1
204
Table 1. General characteristics of the environment
Location French Mediterranean coast
(Languedoc Roussillon)
Watershed name Lez-Mosson
Watershed area (km2 ) 574
Inhabitant/km2 (watershed) 588
Total area (ha) 747
Average depth (m) 0.7
h
Water salinity ( ) 11–36
P input (t/year) 12
Algal growth varies during the year. Seasonal
changes in species composition, biomass and the pro-
duction of macroalgal communities in coastal waters
depend on the interaction of physical, chemical and
biological factors (Thom & Albright, 1990). There
is a close link between the growth and decomposi-
tion of Ulva in particular and phosphate and nitrogen
processes (Viaroli et al., 1991). Phytoplankton appears
in short successive blooms when input nutrients are
high and there is no competition from macrophytes
(Vaulot & Frisoni, 1986).
Sediment plays a major role in the coastal ecosys-
tems because of the lagoons’ shallowness, which
means that it mixes readily with the overlying water.
It can act either as a sink or as a source of phosphate.
Sedimentary phosphate dynamics involve a large num-
ber of mechanisms. Different approaches have been
adopted to study phosphate dynamics in sediment
(López Laseras, 1991): the relationships between the
different descriptors and the established vertical and
horizontal variations, phosphate release rates and the
factors controlling them, and studies of the chemical
fractions of phosphate in sediments.
This study was carried out for the Programme Na-
tional d’Océanographie Cotière (PNOC) with the aim
of understanding the nature of the eutrophication prob-
lem before restorative action is taken. The Méjean
Lagoon (Figure 1), one of the shallow coastal lagoons
bordering the Mediterranean, was chosen.
The study attempts to identify mechanisms ex-
plaining sedimentary phosphate dynamics in terms
of seasonal variations in algal growth. This neces-
sitated simultaneous monitoring of the sedimentary
phosphate fractions and the water conditions.
Materials and methods
Site
The study was carried out using the sediment of the
Méjean Lagoon, a Mediterranean coastal lagoon in the
South of France which receives pollution mainly from
urban waste water. The lagoon has a total surface area
of 747 ha and a mean depth of < 1 m. Variable inputs
h
of fresh water (with rainfall) and sea water cause the
water’s salinity to range from 9 to 31 in the course
of a year. The rate of water renewal is very low in
dry periods (0.01% per day of the lagoon’s volume)
but can reach significantly higher values in periods of
rainfall. The general characteristics of the lagoon are
summarized in Table 1.
Two different sampling points, point 1 in the east-
ern (Pérols) part and point 2 in the western (Méjean)
part, were selected for the removal of sediment sam-
ples. The treated waste water from the Montpellier
region (300 000 habitants) is disposed of in the Mé-
jean area. The Pérols area does not receive any waste
directly from the treatment plant. The hydrodynamics
of the lagoon create little mixing of the water masses
from the Méjean area and the Pérols area. The choice
is the result of the spatial variation study of Gomez
(1996) which showed the existence of two different
parts in this lagoon.
The periods chosen for sediment sampling reflect
the different phases of algal growth.
Sampling techniques
Sediments and pore water were sampled at the same
time as the flux measurements, at both points in April,
June, August and November 1994. The chemical ex-
tractions of phosphate, as well as determination of pH
and EH profiles were carried out in February 1994, in
order to have a winter reference value for the sediment.
Samples from undisturbed sediments were taken
with a manual corer (Rofés & Savary, 1981). The cores
were cut into 5 cm slices (0–5 cm and 5–10 cm), stored
in a plastic flask, kept at 4 ◦ C and treated the same day.
5 cm slices were chosen due to the high proportion
of shells in the sediments which means thinner slices
would be difficult to cut and rather selective. However,
the measurement of pH and EH can be carried out in
situ at a higher resolution by using microelectrodes
placed directly in the sediments. Sediment charac-
teristics were determined after sieving with a 2 mm
sieve.
hy-ja28.tex; 2/09/1998; 19:22; p.2

Lironde
Roubine
Lez
MEJEAN
Mosson
x 1
ARNEL
Rhône-Sète Canal
Palavas
PREVOST
0 1 km
Figure 1. Location of the sampling points in the Méjean Lagoon (France).
The removal of the sediments for extraction of pore
waters was carried out by coring in the absence of
air, in order to avoid any contact with oxygen.These
sediments were then placed in a glove box contain-
ing nitrogen. The porewaters were sampled by the
press method (Reeburgh, 1967), using the pressure of
nitrogen (Mesnage et al., 1993).
Water characteristics
pH, temperature and O2 were measured in the field
directly after sampling. pH was measured with a
WTW pH meter and redox potential with an Ingold
Pt 4800-M5 electrode. Results were corrected for the
standard-H2 electrode (Bühler & Galster, 1980). O2
was measured with a combined oxygen-temperature
WTW92 meter. Soluble reactive phosphate (SRP)
was measured using the standard method (Standard
Methods, 1992). Each macroalgal sample was col-
lected over an area of 0.25 m2 . The samples were
sieved, sorted into species, and the dry weight of
each species was measured after drying at 110 ◦ C for
24 h. Biomass was measured in grams dry weight per
square metre (g dw m−2 ). Phytoplankton was mea-
sured as µg l−1 of chlorophyll a (acetone extraction
and spectrofluorimetry).
205
x Lattes treatment
plant
PEROLS
2
Carnon
GREC
N
Mediterranean Sea
Sediment characteristics
One part of the sediment was freeze-dried to deter-
mine total organic carbon, total nitrogen, and CaCO3
content. Another part was used to determine poros-
ity (Ø = Volwater /Voltotal) and water content (H = water
weight/total weight of wet sediment).
Granulometric distribution in the sediment sam-
ples was determined by laser granulometry according
to Duchaufour’s pedologic nomenclature (1988).
Total phosphate was analyzed, after persulphate
mineralization, as o-P and SRP, according to the
Standard Methods (1992): ammonium molybdate and
potassium antimonyl tartrate react in acid medium
with orthophosphate to form a heteropolyacid-
phosphomolybdic acid, that is reduced to intensely
coloured molybdenum blue by ascorbic acid. Total
iron was determined after Golterman et al. (1978).
Phosphate fractionation was carried out using the
Golterman & Booman (1988)/De Groot & Golter-
man (1990) method, which was appropriate for our
sediments (Mesnage & Picot, 1993). After a water
extraction of weakly adsorbed phosphate (o-P), chelat-
ing agents such as NTA or EDTA were used to extract
iron-bound and calcium-bound phosphate (Fe(OOH)-
P and CaCO3 -P). H2 SO4 0.5M was used to extract
acid-soluble organic phosphate (ASOP) and persul-
hy-ja28.tex; 2/09/1998; 19:22; p.3
206
phate digestion to extract residual organic phosphate
(ROP).
pH profiles and redox potential were measured in
situ, with the electrode introduced into the sediment
every 2 cm in depth.
Statistical analysis
Statistical analysis (Anova, Statview 4) was carried
out to evaluate spatial and temporal variations in phos-
phate concentration in sediments, with three replicates
per measurement.
Abbreviations:
SRP: Soluble Reactive Phosphate in water
o-P: Sediment adsorbed phosphate
Fe(OOH)-P: Ferric hydroxide-bound phosphate
CaCO3 -P: Calcium carbonate-bound phosphate
ASOP: Acid-soluble organic phosphate
ROP: Residual organic phosphate
Tot-P: Total P in sediment
Tot-N: Kjeldhahl N (including NH3)
Tot-C: Total carbon;
Corg: Organic carbon;
Cmineral; mineral carbon.
Results
Water column
Water column characteristics are shown in Table 2.
The temperatures recorded varied between 7.7 ◦ C in
February and 28.2 ◦ C in June. The overlying water
was always sursaturated in dissolved oxygen (mea-
sured near the surface) in the Méjean part and quite
high (at least 77%) in the Pérols part. The highest
values were recorded in August. The redox potential
values were always over 400 mV, in accordance with
the continual presence of oxygen in the water at the
time of sampling. These results are a indication of
continual photosynthesis activity. More frequent mon-
itoring in 1993 showed brief periods of anoxia in early
April, late June and mid-August, always in the Méjean
part (Fillit, 1995).
The mean pH of the water was 8.5 ± 0.3 in the
Pérols part and 8.6 ± 0.4 in the Méjean part. Salin-
h 12.4 and 302.4 g m−2) . Phytoplankton was present in
h
ity varied between 4.2 (Pérols) in November and
27 (Pérols) in June. The lowest values coincided
with periods of heavy rainfall (February, October and
November), while high salinity coincided with periods
of high evaporation (June and August).
The graph of SRP over an annual cycle (Fig-
ure 2) shows higher concentrations over the summer
period (June to October). The lagoon appears to be
divided into two parts, with higher concentrations in
the Méjean than in the Pérols part.
The SRP for the seasonal study varied from close
to zero to 5.16 µM (or ≈ 0 to 160 µg l−1 ) in Pérols
and to 29.68 µM (or ≈ 0 to 920 µg l−1 ) in Méjean.
As in the preceding year, the values were highest in
summer.
The changes in macroalgal and phytoplankton bio-
mass over an annual cycle (Fillit, 1995) show differ-
ences in growth between the two sectors of the lagoon.
The P stored in phytoplankton was calculated using
measured chlorophyl a values, to which the following
conversions were applied: carbon/chlorophyl a = 60
(Strickland, 1960), carbon/nitrogen = 5.5 (Goldman
et Mann, 1980), nitrogen/phosphorus = 16.7 (Harris
et Riley, 1956). The P stored in macroalgae was
calculated using the biomass and phosphate content
measured at each sampling (Fillit, 1995).
– The eastern sector (or Pérols part) is character-
ized by strong phytoplankton growth. The P stored in
the phytoplankton thus represents about 0.5 tonnes of
P throughout the year, except for the June to Septem-
ber period when it is very slight. The macroalgae in
this sector develop from July onwards and their bio-
mass represents, at the maximum, 1.4 tonnes of P in
October. This level then declines.
– The western sector (or Méjean part) is the area
where most macroalgal growth occurred. Only ex-
tremely small amounts of phytoplankton are present
while the macroalgae develop to a massive extent be-
tween April and October. Their biomass reaches a
maximum value of 4.5 tonnes of P.
The western sector can consequently be regarded
as an area of macroalgal growth. The eastern sec-
tor, on the other hand, can be regarded as an area of
phytoplankton growth.
The seasonal survey of algal growth in 1994 (Ta-
ble 2) showed the continual presence of phytoplankton
for the Pérols part (34.8 to 144.7 µg l−1 ) and later
macroalgal growth. The latter were present in minimal
quantities except in November (5.3 g m−2 ). Macroal-
gal growth also occurred later than in the previous year
in the Méjean part, from June to November (between
lower concentrations than in the Pérols part in the four
samples taken at this point (2.5 to 106.5 µg l−1 ). The
hy-ja28.tex; 2/09/1998; 19:22; p.4
 207

Table 2. Water characteristics at time of sampling during 1994 seasonal survey: (×)
below the detection limit, (−) not determined.

Méjean part

Month February April June August November

Temperature (◦ C) 8.5 11.2 27.7 24.6 12.5

O2 (% saturation) 104 95 93 126 77
pH 8.6 – 8.9 8.0 8.9
EH (mV) 418 421 454
Salinity (g l−1 ) 12.5 17.5 23.0 26.0 4.9
Conduct. (mS cm−1 ) – – 36.6 47.5 70.2
o-P (µM) 0.32 3.23 20.00 29.68 ×
Chl a (µg l−1 ) – 6.1 2.5 24.6 107
Algae (g m−2 ) – × 12 302 147

Perols part
Month February April June August November

Temperature (◦ C) 7.7 11.0 28.2 25.5 14.3

O2 (% saturation) 134 111 140 142 123
pH 8.1 8.2 8.7 8.5 8.8
EH (mV) 667 415 440 463
Salinity (g l−1 ) 8.9 18 27 26.5 4.2
Conduct. (mS cm−1 ) – – 39.7 48.5 58.5
o-P (µM) 0.65 1.61 5.16 4.84 ×
Chl a (µg l−1 ) – 98 35 145 60
Algae (g m−2 ) – × × × 5.3
P mg.l-1 (Means and standard deviation)

09-dec 07-feb 08-apr 07-jun 06-aug 05-oct 04-dec 02-feb

Figure 2. Changes in phosphate concentrations (SRP) in the Méjean Lagoon in 1993 (Fillit, 1995). Means and standard deviations in Méjean
and Pérols parths from December to February.

hy-ja28.tex; 2/09/1998; 19:22; p.5

208
Table 3. Sediment characteristics

0–5 cm PEROLS part MEJEAN part

month April June August November mean April June August November mean

Silt < 50 µm – 68 67 68 68 ± 0.6 – 75 77 82 78 ± 4

Fine sand
50–200 µm – 9 9 8 9 ± 0.6 – 7 7 7 7
Coarse sand
200 µm–2 mm – 23 24 24 24 ± 0.6 – 18 16 11 15 ± 4
Water content (%) 64 62 62 61 62 ± 1.3 65 57 62 72 64 ± 6
Porosity (%) 84 78 79 81 80 ± 2.6 81 71 80 87 80 ± 7
CaCO3 (%) – 57 56 56 56 ± 0.6 – 58 53 48 53 ± 5
Tot-P µg g−1 511 ± 25 610 ± 22 580 ± 10 512 ± 44 553 ± 50 701 ± 45 603 ± 24 714 ± 35 901 ± 7 741 ± 110
Norg mg g−1 4.2 4.2 4.2 4.3 4.2 ± 0.05 5.1 5.6 4.8 5.9 5.3 ± 0.5
Tot-C mg g−1 92.3 95.5 94.0 99.6 95 ± 3 93.0 92.5 94.0 98.0 94 ± 2
Cmineral mg g−1 31.3 38.8 45.2 57.5 43 ± 11 27.4 32.2 39.0 45.8 36 ± 8
Corg mg g−1 61.0 56.7 48.8 42.1 52 ± 8 65.6 60.7 55.0 52.2 58 ± 6
Corg/Norg 14.5 13.5 11.6 9.8 12 ± 2 12.9 10.8 11.5 8.9 11 ± 2

5–10 cm PEROLS part MEJEAN part

month April June August November mean April June August November mean
Silt < 50 µm – 72 68 83 74 ± 8 – 73 73 72 73 ± 0.6
Fine sand
50-200 µm – 7 12 3 7±4 – 7 5 6 6±1
Coarse sand
200 µm–2 mm – 21 20 14 18 ± 4 – 20 22 22 21 ± 1
Water content (%) 62 62 60 61 61 ± 1 62 61 58 60 60 ± 2
Porosity (%) 82 79 81 79 80 ± 10 83 77 77 75 78 ± 3
CaCO3 % – 54 55 48 52 ± 4 – 62 59 58 60 ± 2
Tot-P µg g−1 382 ± 4 388 ± 3 350 ± 31 319 ± 24 360 ± 32 543 ± 15 478 ± 31 471 ± 10 494 ± 12 496 ± 32
Norg mg g−1 3.9 3.7 3.9 4.2 3.9 ± 0.2 3.6 4.4 3.5 3.8 3.8 ± 0.4
Tot-C mg g−1 82.0 91.8 92.2 89.4 88 ± 5 92.0 94.0 94.7 98.0 95 ± 2
Cmineral mg g−1 16.6 35.0 39.9 43.2 34 ± 11 36.9 34.6 49.1 55.1 44 ± 10
Corg mg g−1 69.4 56.8 52.3 46.2 56 ± 9 55.1 59.4 38.3 42.9 49 ± 10
Corg/Norg 17.8 15.4 13.4 11.0 14 ± 3 15.3 13.5 11.0 11.3 13 ± 2

values obtained for these four periods showed a trend
similar to that observed the previous year.
Sediment characteristics
The main sediment characteristics are shown in Ta-
ble 3.
The sediment was very fine, with over two thirds
in the < 63 µm fraction (clay and silt). The < 63 µm
fraction represented 79% at Méjean and 68% at Pérols
for the 0–5 cm layer; 74% of the sediment in the 5–
10 cm layer consisted of this fraction at both sampling
points. The water content (62% of wet weight for point
1 and 64% for point 2) and the porosity (80%) of
the sediment in the 0–5 cm layer were fairly constant
throughout the year. Values were of the same order for
the lower layer (5–10 cm). The sediments were cal-
careous (carbonate calcium accounts for 55% of dry
weight on average).
Most of the sediment’s total carbon was organic.
There were higher levels of organic carbon in the Mé-
jean part (65.6 to 52.2 mg g−1 ) than in the Pérols part
(61.0 to 42.1 mg g−1 ). This was due to the anthropic
input (quite considerable in this part of the lagoon)
and the macroalgal biomass that developed at this sam-
pling point. The decrease observed in organic carbon
levels from April to November in both sectors of the
lagoon indicates that a mineralization of organic mat-
ter was enhanced by an increase in temperature. There
was little variation in the levels of organic nitrogen,

hy-ja28.tex; 2/09/1998; 19:22; p.6

 209
Depth (cm) Pérols pH Depth (cm) Pérols EH
6,5 7,0 7,5 8,0 8,5 9,0 -300 -200 -100 0 100 200 300 400 500
10
0

0
-10
-10
February
-20 June
-20 August
February November
April
June -30
-30 August
November

-40 -40

Depth (cm) Méjean pH

6,5 7,0 7,5 8,0 8,5 9,0 Depth (cm) Méjean EH
-300 -200 -100 0 100 200 300 400 500
10

0
0

-10 -10

-20 -20
February
April February
June June
-30 August August
-30
November November

-40
Figure 3. Seasonal variation in the pH of the sediment and the
overlying water for the two sampling points studied.
-40
Figure 4. Seasonal variation in the EH of the sediment and the
overlying water for the two sampling points studied.

which decreased with depth: 4.2 to 5.9 mg g−1 at the

surface and 3.5 to 4.4 mg g−1 at depth. As nitrogen
levels were fairly constant, the Corg/Norg ratio (be-
tween 11 and 17) decreased from April to November
as a result of the reduction in organic carbon.
The pH values (Figure 3) of the sediment were
lower than those of the water in all seasons. Sedi-
ment pH decreased with depth, ranging from 7.7 ± 0.2
(Pérols) and 7.8 ± 0.4 (Méjean) at –2 cm to 7.3 ± 0.2
for both sampling points at –20 cm. The pH profile for
August showed the lowest pH values at both sampling
points.
The redox potential (Figure 4) in the Méjean La-
goon only varied over the first few centimeters of
sediment. After the first two centimeters, it was be-
tween –150 and –200 mV, irrespective of season. The
EH was positive at the surface in February, April and
November, indicating oxic conditions at the interface.
This was not the case in June and August, when the
sediment’s anoxic state made it difficult to measure the
EH at the interface. The lowest values for the year at
–2 cm (in August: –203 mV for the Pérols part and –
276 mV for the Méjean part) were recorded during the
hottest months and provide information about the state
of the interface.
Decomposition of algae led to a major deposition
of organic matter on the sediment, and the carbon
content was always over 40 mg g−1 in the first 10
centimeters. This major deposit of organic matter min-
eralizing in the sediment led to a reduction in pH with
depth and continuing very low redox potential values.

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210
Total phosphate and P fractions
The sediments had a higher P load (Table 3) in the
Méjean (603±24 to 901±7 µg g−1 ) than in the Pérols
part (511 ± 25 to 610 ± 22 µg g−1 ). The values for the
5–10 cm layer were 30 to 40% lower than those at the
surface (471 ± 10 to 543 ± 15 µg g−1 for the Méjean
part and 319 ± 24 to 388 ± 3 µg g−1 for the Pérols
part).
Seasonal changes in total P were quite marked at
the surface (0–5 cm) and different for the two sectors
studied: the maximum values occurred in the Méjean
part in spring and in the Pérols part in winter. The
variations in the 5–10 cm layer are not significant.
Sedimentary phosphate fraction data is given in
Tables 4 (content) and 5 (percentages of total P).
The o-P fraction represented at most 2.7% of total
P. Maximum values were observed in November.
The ferric hydroxide-bound phosphate fraction
(Fe(OOH)-P), which represented between 7 and 31%
of total P, behaved differently in each sector of the
lagoon. The Fe(OOH)-P value of the sediment from
the eastern (Pérols) part of the lagoon was highest in
August (103±2) and lowest in November (67±6) and
February (47 ± 2). In the Méjean part, it was highest
in April (216 ± 14) and lowest during the hot period:
June (112 ± 20) and August (92 ± 2).
The calcium-bound phosphate fraction (CaCO3 -P)
was the largest inorganic fraction. The highest values
were observed in the same season in both parts of the
lagoon, in February for the surface sediments (260 µg
g−1 Pérols, 418 µg g−1 Méjean) and in April for the
sediments of the 5–10 cm layer (188 µg g−1 Pérols,
275 µg g−1 Méjean). These values were not measured
in February. The lowest values appeared at different
seasons in the two sectors of the lagoon. They were
measured in November for the Pérols part (169 µg g−1
at the surface and 106 µg g−1 at 5–10 cm) and in June
for the Méjean part (205 µg g−1 at the surface and
141 µg g−1 at 5–10 cm).
The acid-soluble organic P fraction (ASOP) repre-
sented between 7 and 18% of total sedimentary P. It
was the only fraction whose behaviour was indepen-
dent of the sector concerned: lowest in spring (April)
and highest in summer (June–August) regardless of
the depth.
The measured residual organic phosphate fraction
(ROP) represented between 28 and 56% of total sed-
imentary P. Like the calcium-bound fraction, it was a
major phosphate fraction in the sediment. The high-
est levels for the Pérols part were observed in June
(257 µg g−1 at the surface, 161 µg g−1 at 5–10 cm).
The highest levels for the Méjean part were observed
in November at the surface(393 µg g−1 ) and in June
at 5–10 cm (197 µg g−1 ).
Discussion
Analysis of the exchange situation during the main
algal growth phases
The overall results indicate an influence of the inputs
from the watershed, which were partly responsible for
the high dissolved P values in the Méjean part. The
mean net input (inflow–outflow) was 870±160 kg of P
per month (Burtin 1994 in Fillit 1995). Virtually all of
it came from the tailings of the neighbouring treatment
plants (Lattes – population 6 500 and Montpellier –
population 330 000). Under 10% of inputs are of more
distant origin (upstream Lez, Mosson).
Four phosphate exchange situations were observed
in the west (Méjean part) of the lagoon, where
macroalgal growth is dominant.
In April, when there was an absence of phyto-
plankton and macroalgae in the water column (Ta-
ble 2), conditions were favourable to storage: con-
tinuous inputs from the watershed, no P requirement
by the water column, an oxic layer on the surface of
the sediment and an oxygenated environment. Stor-
age behaviour was in agreement with the experiments
conducted in the laboratory in which all the condi-
tions mentioned were present. The same sediment,
when in suspension (aproximately 5 g l−1 d.w.) in
well-oxygenated water without algae, had adsorbed
300 µg of phosphate addition of 350 µg g−1 (Gomez
et al., 1998). Others sediments show the same behav-
iour (Furumai & Ohgaki, 1988; Istanovics, 1994). On
the other hand, total P values (Table 3) show a major
reduction in the sedimentary load: from 871 ± 20 µg
g−1 in February to 701 ± 45 µg g−1 in April. This
reduction should have led to an increase in the con-
centration of dissolved P in the water column. The
concentration was, however, only 3.23 µm. There are
only two possible explanations for the loss of sedimen-
tary P: either re-suspension of the sediment (and of
the P it contained) or its dilution. Re-suspension can
occur because of wind. In 1993 this phenomenon led
to a noticeable increase in suspended matter, which
exceeded 0.46 g l−1 (Fillit, 1995). Dilution can be
due to high rainfall, which is capable of doubling the
initial depth of the water column. There was a rainy
hy-ja28.tex; 2/09/1998; 19:22; p.8
 211

Table 4. Means of phosphate fractions measured during seasonal survey (phosphate in µg g−1 ) and standard deviations

µg g−1 Month o-P Fe(OOH)-P Fe(OOH) CaCO3-P ASOP ROP Fe/P

February 4 ± 0.5 47 ± 2 891 ± 68 260 ± 15 33 ± 2 151 ± 3 19 ± 0.8

Pérols April 2 ± 0.1 75 ± 10 2054 ± 68 247 ± 17 38 ± 7 148 ± 17 27 ± 2.8
June 2 ± 0.1 85 ± 5 1193 ± 53 175 ± 3 91 ± 1 257 ± 21 14 ± 0.1
0–5 cm August 4 ± 0.3 103 ± 2 1377 ± 54 192 ± 5 74 ± 1 208 ± 18 13 ± 0.2
November 5 ± 0.3 67 ± 6 1153 ± 42 169 ± 8 70 ± 10 201 ± 24 17 ± 0.9

April 2 ± 0.1 59 ± 4 1135 ± 26 188 ± 7 12 ± 3 121 ± 5 19 ± 0.9

Pérols June 1 ± 0.7 44 ± 2 1284 ± 69 111 ± 11 70 ± 2 161 ± 13 30 ± 2.8
August 2 ± 0.1 53 ± 4 677 ± 45 130 ± 5 54 ± 1 111 ± 32 13 ± 0.1
5–10 cm November 7 ± 0.3 41 ± 1 477 ± 12 106 ± 8 49 ± 2 116 ± 16 12 ± 0.3

February 8 ± 0.9 156 ± 10 1721 ± 56 418 ± 3 75 ± 15 214 ± 9 11 ± 0.8

Méjean April 2 ± 0.6 216 ± 14 4058 ± 176 219 ± 24 61 ± 10 203 ± 38 19 ± 0.4
June 1 ± 0.1 112 ± 20 2310 ± 3 205 ± 1 140 ± 22 144 ± 19 21 ± 3.7
0–5 cm August 8 ± 0.2 92 ± 2 1693 ± 186 216 ± 2 121 ± 19 277 ± 13 18 ± 2.3
November 35 ± 8 116 ± 5 1761 ± 131 256 ± 5 101 ± 15 393 ± 2 15 ± 0.5

April 1 ± 0.3 101 ± 1 3151 ± 1 275 ± 213 20 ± 14 146 ± 4 31 ± 27

Méjean June 3 ± 0.2 45 ± 1 516 ± 18 141 ± 24 92 ± 33 197 ± 21 11 ± 0.1
August 3 ± 0.3 78 ± 1 1094 ± 29 165 ± 12 96 ± 3 128 ± 20 14 ± 0.5
5–10 cm November 8 ± 0.1 74 ± 1 1103 ± 28 195 ± 9 62 ± 9 155 ± 17 15 ± 0.3

Table 5. Percentages of phosphate fractions measured during seasonal survey

% Month o-P Fe(OOH) CaCO3-P ASOP ROP

February 0.8 10 52 7 31
Pérols April 0.3 15 48 7 29
June 0.2 10 20 10 42
0–5 cm August 0.5 13 24 9 36
November 0.7 9 24 10 56

April 0.4 16 49 3 32
Pérols June 0.2 8 20 13 42
August 0.4 11 29 12 31
5–10 cm November 1.5 10 24 11 36

February 1.0 18 48 9 25
Méjean April 0.2 31 31 9 29
June 0.2 19 34 23 24
0–5 cm August 1.1 13 30 17 39
November 4.0 13 28 11 44

April 0.2 19 51 4 27
Méjean June 0.4 7 21 13 41
August 0.5 13 28 16 27
5–10 cm November 1.2 11 30 10 31

hy-ja28.tex; 2/09/1998; 19:22; p.9

212
period in February 1994 which might have been re-
sponsible for the loss of considerable quantities of P
from the lagoon. During a similar rainfall, 33% of the
total water volume of the lagoon was renewed in less
than six days. This would explain why concentration
increased by only 2.90 µm during the period under
consideration whereas it should have been higher in
view of the inputs (from the watershed and release by
the sediment).
In June, the algal biomass in the water was low
(2.5 µg l−1 of Chla and 12.4 g m−2 of macroalgal
biomass) and the concentration of P in the water very
high (20 µm). The dissolved P came from the sed-
iment, and the load diminished between April and
June from 701 ± 45 to 603 ± 24 µg g−1 , which was
statistically significant. The increase in temperature
compared to that of April was very considerable (from
11.2 to 27.7 ◦ C). Biological activity should therefore
have been stimulated both in the water, which had
a pH of 8.9, and in the sediment. Phosphate release
can be caused by mineralization of the organic matter
by bacterial activity, as Hasson (1989) observed un-
der anaerobic conditions. This release was increased
by the establishment of reduced conditions at the
water–sediment interface.
In August 1994, as in the previous year, algal
growth was very heavy (302 g m−2 of macroalgal
biomass) with much sunshine and high temperatures.
Despite the increased sedimentary load, P release
appears to have continued after June, as the concentra-
tion of dissolved P in the water (29.68 µm) increased.
There was a simultaneous growth of macroalgae and
degradation of part of the biomass during this period
of algal growth. The observed increase in the sedi-
mentary load was caused by the accumulation of dead
biomass in the sediment.
In November, there was a reduction in macroal-
gal biomass compared to the August level, but it was
still considerable (147 g m−2 of macroalgal biomass).
Phytoplankton bloom was observed during this sam-
pling period (106 µg l−1 of Chla). The heavy algal
presence was responsible for the disappearance of P
in the water (value below the detection limit). The
fall in temperature from 24.6 to 12.5 ◦ C slowed down
biological activity in the sediment and enabled the sed-
iment surface to return to oxic conditions, favouring
storage. The load of sedimentary P increased again
between August and November, due to the deposition
of decomposing macroalgae.
The annual pattern of changes was completely
different in the Pérols part.
The algal population consisted solely of phyto-
plankton growth (Table 2), with macroalgae present
only when the November samples were taken and
even then in small quantities (5.3 g m−2 ). The rate of
phytoplankton renewal was much more rapid that for
macroalgae, which explains its appearance in the form
of brief, successive blooms. Fillit (1995) had shown
a continuous existence of phytoplankton in this part
of the lagoon in 1993, with high chlorophyll a values
in winter and low ones in the summer (September).
In 1994 phytoplankton growth occupied the stretch of
water, with a maximum level in August, in the absence
of competition from macroalgae.
Seasonal changes in the load of sedimentary P
were smaller and the reverse of those observed in
the Méjean part. Only the increase observed between
April and June and the reduction between August and
November are significant. The P accumulating in the
sediment between April and June could not have come
from the macroalgae, which were absent in this sector
of the lagoon, or from the inputs from the watershed,
which remained fairly constant over time. The temper-
ature increase led to a change to reduced conditions in
the sediment from July onwards. The resulting release
is indicated by the reduction in the sedimentary load
between August and November.
Sediment
The high primary production of the Méjean Lagoon
led to major inputs of organic matter in the sediment,
which were reflected in low redox values, in the order
of –150 mV (Figure 4). The results obtained for the
EH profiles in this eutrophic environment are similar
to those found in summer in the oyster-farming area
of the Thau Lagoon (Mesnage et al., 1994; Mesnage
& Picot, 1995). Several authors place the iron oxi-
doreduction, controlling the availability of some of the
phosphate trapped by the sediment within this range of
observed redox values (Whitfield, 1969; Bagander &
Niemisto, 1978).
The P fractions in the Méjean Lagoon showed
a mobility capable of creating fluxes at the water–
sediment interface as a function of the environmental
variates.
Macroalgal growth and mortality were also in-
volved in the phosphate exchanges in the Méjean
part.
The presence of an oxic layer on the sediment
surface favoured adsorption of the P by the ferric
hydroxide-bound fraction. Adsorption was confirmed
hy-ja28.tex; 2/09/1998; 19:22; p.10

by the increase in Fe(OOH)-P between February and
April. The increase in this fraction under quite high
redox conditions agrees with the observations of nu-
merous authors, both in the field (Detenbeck & Bre-
zonik, 1991; Nair & Balchand, 1993), in laboratory
experiments (Furumai & Ohgaki, 1989) and in our
own experiments (Gomez et al., 1998). In June the
temperature increase activated bacterial degradation
of organic matter (Organic carbon decrease, Table 3).
This phenomenon caused a fall in EH (–150 mV) over
the first five centimeters of sediment and provoked
the release of phosphate and iron (1.7 mg g−1 of Fe)
from the Fe(OOH)-P fraction. At the same time, the
pH on the surface sediment increased ( 0.8 units).
Numerous authors, including Boström et al. (1988)
and Seitzinger (1991), have shown that an increase
in pH also reduces the adsorption capacity of ferric
hydroxides.
The calcium-bound phosphate fraction is sensitive
to pH variations in the surface layer: it declined be-
tween February and April because of a 1-unit fall in
pH. Its mobilization in the water explains the reduction
in the load of sedimentary P observed between Feb-
ruary and April. CaCO3 -P increased again between
August and November at the same time as pH. Vari-
ations in this fraction are significant only in the two
situations mentioned. Variations in pH do not produce
the same results systematically as they are combined
with variations in other variates (Jensen & Andersen,
1992). In addition, laboratory experiments with cal-
cium salts confirm that the latter play an active part in
P retention when there is a high pH. Stumm & Mor-
gan (1981) show the formation of hydroxyapatite at
high pH and other authors suggest increased precip-
itation of P and calcite when the pH rises (Ishikawa
& Ishikuni, 1981; Lopez & Morgui, 1992). Precipita-
tion of P with calcite occurs very rapidly (10 to 100
minutes), and is followed by a slow phase (Otsuki
& Wetzel, 1972; House & Donalson, 1986; House
et al., 1986; Suzuki et al., 1986). Work by Martens
(1970) and Guldbransen et al. (1984) has shown, how-
ever, that precipitation with calcium is slower, in the
presence of magnesium, in marine environments.
The organic phosphate (ROP) in the surface sed-
iment increased after degradation of the lower layers
of macroalgae, which were deposited on the sediment
from August onwards. The organic carbon measured
during this period decreased, despite the deposition
of macroalgal debris. This indicates a more rapid de-
composition of carbon compounds than of phosphate
compounds.
213
The difference in concentration observed in No-
vember between the pore water (Table 4) and the
overlying water (values below the detection limit) may
be responsible for a diffusive P flux at the interface
(Enell & Löfgren, 1988). The flux will tend to equi-
librate the two concentrations at an equilibrium value
such as has been measured by McCallister & Logan
(1978), Froelich (1988) and Oliya & Reddy (1993).
The consumption of P by the phytoplankton accounts
for the continuing gradient at the interface.
The load of total P increased in June in the Pérols
part (Table 3) because of the organic fractions. The
origin of the organic P cannot be explained on the
basis of the data collected for this sector (absence of
macroalgal bloom). A transfer of dissolved P from
the Méjean sector is conceivable, given the consider-
able difference in concentration (20 µM in Méjean,
5.16 µM in Pérols).
The organic fractions declined between June and
August because of bacterial degradation, which in-
creased with temperature (Table 2). In aerobic envi-
ronments bacterial activity is responsible for an in-
creased rate of P release (Sinke & Cappenberg, 1988).
Release may occur by oxidation of the organic forms
of P or by dissolution of the CaCO3 -P following a
drop in pH. In anaerobic environments, bacteria are re-
garded as catalysts for iron hydroxide reduction which
results in the release of iron-bound P, in other words as
an indirect source of P through the use and release of
internal P accumulated in aerobic conditions (DeMon-
tigny et al., 1993). In the present case, ferric hydroxide
reduction cannot be considered as the redox potential
remained unchanged (∼ −180 mV). In addition, the
Fe(OOH)-P fraction increased, even if only slightly.
It is therefore likely that mineralization of the organic
fractions is a relevant factor in explaining the release.
The ASOP fraction is considered to be an available
P fraction (Golterman & Booman, 1988); it is there-
fore logical that this fraction should be mobilized.
The same is also true, however, for one part of the
ROP fraction, whereas it had initially been regarded as
non-degradable. Research by Mesnage (1994) shows,
however, that this fraction can be involved in phos-
phate exchanges. De Groot & Golterman (1993) show
the presence of phytate in this fraction on paddy field
sediments.
The inorganic fraction was involved when the
degradation of organic matter creates strong reduc-
ing conditions in the surface sediments during August
(Figure 4). Between the measurement in August, when
the sediment did not have an oxic layer at the surface,
hy-ja28.tex; 2/09/1998; 19:22; p.11
214
and November, the load of sedimentary P declined
because of the Fe(OOH)-P and CaCO3 -P fractions.
The iron-bound fraction was mobilized at the same
time as the ferric hydroxides in reducing conditions,
which corresponds to the traditional theory of P re-
lease by iron reduction (Mortimer, 1941, 1942; Patrick
& Khalid, 1974; Sundby et al., 1992).
The strong phytoplankton presence maintains high
dissolved oxygen concentrations and a high pH in the
water in winter. Because of these two conditions, both
phosphate adsorption by iron and phosphate precipita-
tion with calcium are possible. The sediment load is
replenished by sedimentation of inorganic P.
Conclusion
The Méjean Lagoon is a coastal lagoon subject to
severe eutrophication, reflected in a proliferation of
macroalgae and phytoplankton over the course of the
year. The seasonal cycle of the dissolved P in this
shallow lagoon is determined not by inputs from the
watershed, which are relatively regular, but by factors
internal to the lagoon itself: the biological cycle of the
macroalgae, and release from the sediment.
A climatic factor, temperature, regulates the be-
haviour of the sedimentary phosphate in the Méjean
Lagoon. The high temperatures of the hottest months
in the region trigger the release flux following degra-
dation of the sediment’s organic matter. The Novem-
ber fall in temperature slows down biological activity
in the sediment and, by the same token, phosphate
release.
Replenishment of the sediment’s phosphate stock
starts when macroalgal mortality reaches major pro-
portions and continues throughout winter. Macroalgal
mortality also has the effect of prolonging the phe-
nomenon of sedimentary phosphate release by decom-
position of the deposited algal biomass.
In the absence of macroalgae in the sector charac-
terized by phytoplankton growth, there is no length-
ening of the autumn release and the sedimentary load
of organic phosphate can hardly be replenished in au-
tumn. The phosphate load is replenished in winter by
an increase in the inorganic fractions caused by the
oxic conditions prevailing in the environment and by a
high level of phytoplankton activity in this sector.
Macroalgal growth appears to be a phenomenon
unconnected with phosphate release from the sedi-
ment. The inputs from the watershed in fact give rise
to sufficiently high concentrations of dissolved P for
algal growth. Fillit (1995) has shown that in the Mé-
jean Lagoon this element is never limiting macroalgal
growth and concludes that there must be limitation by
light and temperature.
The phosphate released by the sediment is not es-
sential to phytoplankton growth, as the highest chloro-
phyll a values are recorded in winter, when there is no
release. The phytoplankton develops on the strength of
the inputs from the watershed and takes advantage of
the phosphate supplied by the sediment in periods of
release.
Phosphate inputs arriving in the Méjean Lagoon
come mainly from the tailings of the treatment plants.
The first step towards the rehabilitation of the lagoon
is for inputs from the watershed to be reduced or even
stopped.
If inputs stopped completely, the supply of phos-
phate for the water column and hence for macroal-
gal growth would depend for a long time on the
release of P by the sediment. As the amount of phos-
phate released is greater than the amount necessary
for macroalgal growth, phosphate would not be a
growth-restricting factor.
It is likely that the same amount of biomass will be
produced as at present in the area in which macroal-
gae are the dominant feature. The deposition of this
biomass on the sediment would replenish the sedimen-
tary load of P. The system would lose phosphate only
through the process of water renewal, which allows
dissolved P to depart.
A relatively long period thus appears necessary
before the phosphate starts to restrict algal growth
and, by the same token, before any hypothetical
rehabilitation of the lagoon can be effective.
Acknowledgements
This research was carried out with the financial sup-
port of the Programme National d’Océanographie
Côtière.
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