Biological and Physical Events Involved

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International Journal of Salt Lake Research 4: 335-347, 1996.

(~) 1996 Kluwer Academic Publishers. Printed in the Netherlands.

Biological and physical events involved in the origin,


effects, and control of organic matter in solar saltworks

J O S E P H S. DAVIS t and M A R I O G I O R D A N O 2
1Department of Botany, University of Florida, 209 Carr Hall Gainesville, FL 32611, USA;
2Facolt6 di Scienze, Universit6 di Ancona, 60131 Ancona, Italy

Abstract. Aspects of communities and events in the concentrating ponds (S.G. 1.130 to 1.214)
and salt crystallizing ponds (S.G. 1.215 to 1.264) of solar saltworks pertinent to salt manu-
facture are described. Communities that aid salt manufacture enable continuous and efficient
production of high quality salt at a saltworks' design capacity, and they provide important
controls on levels of organic matter in the brine. Fluctuating salinities, high concentrations of
nutrients, and petroleum products are disturbances that cause Aphanothece halophytica and
Dunaliella salina to release excessive quantities of organic matter, and that suppress or cause
death to nutrient stripping organisms. Disturbances result in decreased quality and quantity
of salt and increased costs for salt harvest, washing, and pond upkeep. Organic matter can
be controlled by management techniques that keep nutrient stripping communities at proper
levels and maintain a narrow and unchanging range of salinities in each pond, by constructing
pond dikes able to withstand wind and water erosion, and by preventing spills of petroleum
products in the ponds.
Key words: Aphanothece halophytica, A rtemia, biological management, Dunaliella, halobac-
teria, hypersalinity, Myxobaktron salinum, organic release, sodium chloride, solar saltworks,
Synechococcus sp.

Introduction

One third o f the worldwide sodium chloride (salt) production occurs in solar
saltworks (salinas, saltfields) by solar evaporation of seawater or inland brines.
Solar saltworks are one to several series o f interconnected concentrating ponds
( 4 0 - 8 0 cm depth; S . G ) 1.026 to 1.214 with seawater intake; S.G. 1.1202 to
S.G. 1.214 with inland brine intake) and crystallizing ponds ( 3 0 - 5 0 cm depth;
S.G. 1.215 to 1.264) in which the concentration o f sodium chloride increases
as the water or brine evaporates and flows downstream. After becoming
saturated with salt in the concentrating ponds, the brine is pumped or flows
into crystallizer ponds (crystallizers) where evaporation continues and sodium
chloride precipitates on the bottom. The supernatant liquid (bittern), rich in
c o m p o u n d s o f magnesium, potassium, and sulfate, is removed and the sodium
chloride is harvested with mechanical equipment. The salt is then washed and
stored for a time on a stockpile, where after significant amounts of water and
336

contaminants drain away, the sodium chloride is about 99.7 per cent pure on
a dry weight basis.
The highly competitive solar salt industry and the low market value of
sodium chloride (about US$20.00 per ton) require efficient production of
high quality salt at a saltwork's design capacity. Efficient salinas continuously
produce about 1000 metric tons of salt per ha of crystallizer surface per year,
have stable operating and production costs, allow little or no brine leakage
from the ponds, and have losses of salt in the washing process not exceeding
12 per cent. Quality of salt ready for market is determined by characteristics
of the salt crystals (hollow or solid, and size), and by concentrations of
contaminants (calcium, magnesium, sulfate, and insoluble matter) in the salt.
Premium prices are paid for large solid crystals (Butts, 1977) and for salt
with contaminants not exceeding 0.03 to 0.05 per cent calcium, 0.02 to 0.04
per cent magnesium, 0.11 to 0.16 per cent sulfate, and 0.01 to 0.02 per cent
insoluble matter (Davis, 1990).
A group of living organisms, herein designated a biological system,
inevitably develops in the ponds of every solar saltworks. A biological sys-
tem that aids salt production maintains a variety of species that reproduce
and release organic matter at levels to enable efficient production of high
quality salt at design capacity. A biological system that harms salt production
consists of few species and accumulates excessive amounts of organic matter
that prevents efficient production of high quality salt at design capacity.
Deleterious effects of organic matter for salt production in ponds of low
(S.G. 1.026 to 1.075) and intermediate salinity (S.G. 1.075 to 1.125) are
well documented, and management procedures to control these substances
are widely practised (e.g., Davis, 1980, 1990, 1993; DeMedeiros Rocha and
Camara, 1993; Sammy, 1983). However, detrimental aspects of organic matter
that originates in the highly saline concentrating ponds (S.G. 1.120 to 1.214)
and crystallizers have received little attention. This paper, based on worldwide
observations and successful management techniques by one of us (J.S.D.) at
many seasonal and continuously-operated solar saltworks, and on laboratory
research of both of us and others, reviews aspects of physical and biological
events in ponds at S.G. 1.120 to 1.264 pertinent to salt production, reports
on the origin and effects of excessive nutrients, and describes methods that
have been used to control excessive quantities of organic matter in the highly
saline concentrating ponds and crystallizers of commercial solar saltworks.

Physical events pertinent to salt production

Seasonally operated solar saltworks in dry temperate climates that have well-
defined rainy and dry months (e.g., on the shores of the Black Sea, Mediter-
337

ranean Sea, San Francisco Bay, South China Sea) maintain a salinity gradient
throughout the ponds only during summer months, but continuously operat-
ed salinas in dry tropics (e.g., Bahamas, Brazil, Mexico, Western Australia)
maintain the gradient throughout the year. Unidirectional (downstream) brine
flow through the ponds may be continuous, or may proceed from pond to
pond after appropriate salinities are reached.
Brine of the highly saline concentrating ponds is saturated with calcium
carbonate and precipitates calcium sulfate (gypsum). Gypsum deposition at
S.G. 1.120 appears as small, widely scattered calcium sulfate crystals on
the surface of the benthic community. Precipitation downstream progresses
to thin, continuous sheets which become thick (often 25 mm to more than
100 mm) and firm near S.G. 1.180. As the brine approaches saturation with
sodium chloride, the quantity of gypsum diminishes. In crystallizers, gyp-
sum precipitates until about S.G. 1.230 is reached, but magnesium sulfate is
deposited at higher densities. Daytime temperatures of the brine of highly
saline concentrating ponds and crystallizers are well above ambient, the brine
pH varies from 7.4 to 7.9, and the dissolved organic content ranges from
50 to 200 mg L -1 (Sammy, 1983, original data). Reported concentrations
of combined nitrogen and phosphate, mainly from filtered samples, are near
15 #M for NH +, 10 #M for NO 3, and 1 #M for phosphate (Britten and John-
son, 1987; Coleman and White, 1993; Davis, 1978; Javor, 1983; Landry and
Jaccard, 1982; Nixon, 1970; Rahaman et al., 1993b; Sammy, 1983; original
data).
In seasonal salinas, salt is precipitated on crystallizer floors of compacted
clay, marl, mud, or sand. At harvest time, the 5 to 15 cm layer of salt that
has been deposited and part of the floor materials are picked up by harvest
equipment, loaded onto vehicles, and transported to a salt washing facility.
Jets of seawater or brine under high pressure, and screens of appropriate mesh
sizes significantly decrease contaminants in the salt. In continuously operated
saltfields with seawater intake, or of saltworks with inland brine intake (e.g.,
La Pampa, Argentina, Lake MacLeod, Australia, and Sua Pan, Botswana), a
salt layer (10-15 cm or greater) deposited on the natural floors is not harvested
but becomes a 'salt floor' above which the harvestable salt is precipitated. The
'salt floor' allows a harvest relatively free of natural bottom contaminants,
and provides a firm base that supports harvest and grading machinery.

Biological events pertinent to salt production

Benthic and planktonic communities compose the biological system. Biodi-


versity is highest along S.G. 1.120 to 1.140, but the variety of species begin
to decrease at higher salinities where gypsum precipitates thickly on pond
338

Phosphate
mg L ' I
:Brine: . . . . . .
0.06
-~

3 . 0 mm " Layer I 0.7


1 . 5 mm 9 Loyer 2 1.3
2 . 5 mm i III I I
: Layer 3 3.0

1 9 . 0 0 mm 9 Layer 4 ~30

Fig. 1. Sectionof a bottom communityfrom a dry tropics pond at S.G. 1.120. Orthophosphate
in the brine and in each layer was measuredby a modificationof the method of Helyar and
Brown (1976).

floors. At saturation with sodium chloride, the few species that exist are often
present in high concentrations (Oren and Dubinsky, 1994).
In areas of the floors of highly saline ponds without gypsum or where gyp-
sum is present only in thin, isolated patches of crystals, the benthic community
forms a firm, multilayered structure 20 to 50 mm thick or greater. Figure 1 is
an example of a bottom community at S.G. 1.120 from a five-year-old pond.
The yellow, orange, or brown uppermost layer consists of blue-green algae
(Aphanothece halophytica, Oscillatoria sp., Spirulina sp., Xenococcus sp.),
green algae (Dunaliella salina), and aerobic halobacteria (mainly Halobac-
terium spp.), but diatoms, ciliates, flagellates, and other blue-green algae and
bacteria are also present. The second layer contains organisms similar to the
uppermost, but the algae are bright green. The third layer, bright red, is main-
ly constituted by the purple bacteria Chromatium sp. The lowermost layer
is gray to black and consists of Beggiatoa spp. and other anaerobic bacte-
ria, organic matter, and sulfide-rich muds. In areas of the floors completely
covered with gypsum, the benthic community has the aspect of a green lay-
er including Aphanothece halophytica, Dunaliella salina, Oscillatoria sp.,
flagellates, ciliates, and bacteria that grow among the bases of the gypsum
crystals, in the underlying organic matter and in the natural bottom soils. The
benthic community serves to strip and sequester minerals from the overlying
brine (Davis, 1990, 1993), control brine leakage (Davis, 1978; Jones et al.,
1981), and decrease the rate of gypsum accumulation (Geisler, 1982).
Worldwide, the planktonic community of the highly saline concentrat-
ing ponds consists mainly of Artemia spp. (brine shrimp), DunalielIa sali-
na, Ephydra (brine fly) larvae, chironomid larvae, and species of aerobic
halobacteria, but Aphanothece halophytica, and species of other algae, bac-
teria, flagellates and ciliates are also present. In the dry tropics salinas of
339

Fig. 2. PlanktonicSynechococcussp. (sphericalto ovalcells, 10-20#m dia.)andMyxobaktron


salinum (elongatecells,4-5 #m dia.).

the Bahamas, Caribbean Sea, Israel, Mexico, and South America, the plank-
ton is supplemented by Myxobaktron salinum (Dactylococcopsis sp.) and/or
Synechococcus sp. (Golubic, 1980; original data), which appear abruptly near
the beginning of the high salinity range (S.G. 1.120) and disappear at satura-
tion with sodium chloride (Fig. 2). These highly vacuolate, non-mucilagenous
blue-green algae are yellow-orange, thrive in brines restricted to narrow salini-
ty ranges (Golubic, 1980; original data), adapt to high light intensities, and are
often present in concentrations that color the brine dark brown (original data;
Van Rijn and Cohen, 1983). The planktonic community serves to increase
solar energy absorption, elevate brine temperatures and evaporation (Davis,
1993; Jones et al., 1981, Tackaert and Sorgeloos, 1993), and strip important
quantities of combined nitrogen and phosphate from the brine (original data).
Large brine shrimp populations able to strip significant quantities of nutri-
ents from the brine are essential to salt production (Davis, 1980, 1990, 1993).
These animals thrive in the highly saline concentrating ponds, but their num-
bers and activities decline sharply in the crystallizers. Artemia ingest plank-
ton, organic and inorganic particulates in the brine, oxidize much of the
organic matter they ingest, and release waste products in membrane-bounded
fecal pellets which become incorporated into the benthic communities. Brine
shrimp ingest and utilize Dunaliella salina, Myxobaktron salinum, and Syne-
chococcus sp. (Davis, 1978; original data; Walsby et al., 1983). Mineral
340

stripping by thriving populations of Dunaliella salina, Myxobaktron salinum,


and Synechococcus sp., and voracious feeding by Artemia constitute effective
controls on organic matter in the brine. Artemia may ingest but do not utilize
Aphanothece halophytica (DeMedeiros Rocha, 1993; Gibor, 1956; original
data); the brine shrimp population will decline or die if no other food is
available.
The benthic community of crystallizers (1 to 3 mm thick or more), located
between the salt and the natural floor materials, or the salt floor and the native
bottom substances, is organized into an upper green layer ofDunaliella salina,
aerobic and anaerobic halobacteria, and OsciUatoria sp., and a lower gray-
black layer of anaerobic bacteria and organic matter. The benthic community
contributes to the control of brine leakage and aids degradation of organic
particulates.
The brine of crystallizers may be clear, or it may be colored pink or orange
to red by planktonic organisms imported from the highly saline concentrat-
ing ponds, and/or by algae and bacteria originating within the crystallizers.
Clear brine is the consequence of low concentrations of suspended pigmented
organisms. Orange to red colors are caused by DunalieUa salina cells, rich
in beta-carotene; pink colors are determined by pigmented aerobic Halobac-
terium and other aerobic halobacteria, whose cells have a high content of
bacterioruberin (Oren and Dubinsky, 1994). Dense populations of aerobic
halobacteria not only serve to increase evaporation, but oxidize significant
quantities of dissolved and particulate organic matter, and organisms (includ-
ing Artemia) in the brine (original data).

Origin, effects and control of excessive organic matter

Origin of excessive organic matter

Excessive organic matter is the consequence of disturbances that allow organ-


isms usually restrained by low concentrations of nutrients and high salinity to
reproduce at high rates and release large amounts of organic substances, and
that suppress or eliminate other organisms (e.g., Binford et al., 1983; original
data). Disturbances may develop slowly after excessive nutrients (combined
nitrogen, phosphate, and organic matter) enter ponds over a period of time at
concentrations greater than needed to maintain communities at levels asso-
ciated with efficient production of high quality salt at design capacity, and
after steep increases in brine salinity. Disturbances may develop quickly after
severe brine dilution from rainfall and breach of dikes, and from release of
petroleum products in the ponds. Simultaneous occurrences of both types of
341

disturbances result in release of massive quantities of organic matter (original


data).
Most released organic substances are from Aphanothece halophytica and
Dunaliella salina that thrive after disturbances caused by excessive nutrients
and severe brine dilution (original data). Both organisms utilize a variety of
nitrogen sources and tolerate a wide range of salinities (Berland et al., 1989;
Brock, 1976; Davis, 1993; Ginzburg et al., 1990; Giordano et al., 1994; Loe-
blich, 1972; Takabe, 1988; Tindall et al., 1978; original data). Aphanothece
halophytica populations release mucilaginous lipopolysaccharides (Jones and
Yopp, 1979) that form hyaline sheaths around individual cells and groups of
ceils. Dunaliella salina populations release small amounts of organic mole-
cules (Giordano et al., 1994; see also Huntsman, 1972), enzymes (Giordano,
1992), and glycoproteins (Chardard, 1990). In salinas whose intake is inland
brine nearly saturated with sodium chloride, Dunaliella salina is the chief
producer of organic matter (original data; Sammy, 1983).
In brines saturated with sodium chloride, severe dilution, and release of
petroleum products from pumps, ruptured pipelines, grading equipment and
harvest machinery may burst or decimate aerobic halobacteria and other
resident biota (e.g., Mohr and Larsen, 1963; original data) and result in steep
increases in organic matter in the brine. In laboratory experiments, Ward and
Brock (1978) have demonstrated that metabolic activities of organisms in
brine saturated with sodium chloride are severely depressed in the presence
of certain petroleum products.

Effects of excessive organic matter

(a) Destruction of nutrient-stripping communities


Disturbances that result in excessive reproduction and mucilage release by
benthic Aphanothece halophytica may cause large sections of the bottom
community (often 25 to 50 mm thick and 100 to 500 mm 2 or greater) to rise
to the surface, float downstream, and slowly dissolve (Davis, 1990). The float-
ing rafts cause anoxic conditions during darkness, and dramatically decrease
the Artemia, Myxobaktron salinum, and Synechococcus sp. populations, and
almost all other species in the brine (original data). Upon reaching brine sat-
urated with sodium chloride, most organisms in the floating sections perish,
and the mucilaginous substances completely dissolve. The brine then increas-
es in viscosity (Coleman and White, 1993), becomes brown or black (original
data; Rahaman etal., 1993a, b), and destroys the aerobic halobacteria (original
data).
Steep increases in brine salinity may result in gypsum deposition that
completely covers benthic communities throughout the highly saline concen-
342

b q
t
4 : i

i
C
C
q
q
I

Fig. 3. Progressive deterioration of salt floors in crystallizers with increasing destruction


proceeding at the perimeters (upper row), or progressing from one side to the other (lower
row). Shaded areas denote floorsunable to supportheavy machinery.

trating ponds. The mineral stripping activities of the communities are then
severely diminished.

(b) Decrease in surface areas and volumes


Disturbances resulting in fast accretion of benthic communities that remain
on pond bottoms decrease surface areas and volumes (original data). These
ponds then deliver less brine to the crystallizers. Decreased volumes of ponds
result in increased brine dilution when rainfall occurs.

(c) Deterioration of salt crystal characteristics and destruction of salt floor


integrity
The distribution of crystal sizes and crystal characteristics in the salt profile
(harvestible salt and salt floor) changes as organic matter accumulates. Under
these conditions the larger solid crystals become restricted to the upper part
of the deposit, and the smaller to the lower portion; continued accumulation
results in hollow crystals, and/or small crystals (all less than 1 mm on a side)
with brown to black inclusions in the whole salt layer.
Figure 3 depicts progressive deterioration of salt floors in crystallizers that
have accumulated excessive organic matter. Cross-hatched areas represent
portions of a salt floor composed of small, separated crystals unable to support
harvest equipment. Deterioration may begin at the periphery of a crystallizer
and progress toward the center (upper row), or the damage may originate at
one end of the pond and continue toward the other until the floor is entirely
343

destroyed. Harvest then becomes difficult or impossible, and the ponds may
be abandoned or require expensive corrective action. Sarig et al. (1975) and
Sarig and Tartakovsky (1975) have reported that organic matter alters crystal
shape and prevents close coherence of crystals.

(d) Contaminant increases in salt


Excessive organic matter complexed with calcium (Barcelona and Atwood,
1978) in highly saline concentrating ponds contributes to the deposition of this
mineral in crystallizers (Burnard and Tyler, 1993; Davis, 1990; original data).
Magnesium compounds bind to organic matter that coats salt crystals pre-
cipitated from mucilaginous brine (Davis, 1990); magnesium contamination
levels rise as the size of the sodium chloride crystals decreases and the surface
area of the crystals increases (Davis, 1990). Hollow and misshapen crystals
(hoppers) that develop from mucilage-laden brines (Baha AI-Deen and Baha
A1-Deen, 1973; Davis, 1990) retain liquid contaminants whose composition
is similar to the solution from which the crystals were precipitated (Dubessy
et al., 1983; Geisler, 1982).

(e) Cost increases in salt transport and salt washing


Liquid drainage from vehicles loaded with freshly harvested small and hollow
salt crystals precipitated from mucilagenous brine may significantly increase
transport costs from crystallizers to the salt washing facility, and may damage
the transport roads. Equipment required to wash this salt to acceptable con-
taminant levels is more complex, consumes more energy, requires increased
maintenance, and results in greater losses of salt than machinery needed to
wash large, solid crystals. Use of caustic soda and centrifuges to remove
magnesium from the crystals may add nearly one US dollar per ton to the
cost of salt production.

Control of excessive organic matter

Appropriate salina design tailored to climate, demographics, and ecology of


the area that permits biological management by timely small energy subsi-
dies prevent, minimize, or allow fast correction of the effects of disturbances
(Davis, 1993). However, in the highly saline concentrating ponds and crys-
tallizers of any saltfield, organic matter can be controlled by management
practices that maintain biodiversity, restrain reproduction and organic release
by Aphanothece halophytica and Dunaliella salina, permit communities of
microorganisms and brine shrimp efficiently to strip and oxidize nutrients
from the brine (Coleman and White, 1993; Davis, 1993), and reduce the
effects of disturbances on communities.
344

Of control practices for the highly saline concentrating ponds, maintenance


of the salinity of each pond or in each segment of a pond in a narrow and
specified range is the most important (e.g., Bremer, 1983; Davis, 1990). This
control measure can be implemented by adjusting flow rates during periods
of low and high evaporation and after rainfall, increasing the brine depths
to lessen dilution by rainfall, using decanters to drain rainwater from pond
surfaces (Davis, 1990), counteracting expected rainfall by allowing salt to
precipitate in concentrating ponds usually saturated with sodium chloride,
and providing spare pumps at pump stations.
Additional control measures effective for limiting nutrients within the
highly saline concentrating ponds include use of armored, appropriately ori-
ented dikes that partially traverse large ponds to improve brine circulation
over mineral-stripping bottom communities, prevent brine short circuits, and
decrease destruction of benthic communities by wind and waves, and con-
struction of dikes at pond peripheries able to withstand erosion from wind
and waves, and to prevent land runoff water from entering ponds. Other man-
agement practices include adjustment of brine depths to allow light to reach
benthic communities, introduction or reintroduction of Myxobaktron salinum
and Synechococcus sp., and careful management of the Artemia population.
Procedures to maximize nutrient stripping by brine shrimp include, in
addition to the above control measures, adjustment of brine depths to permit
appropriate temperatures for the animals (Tackaert and Sorgeloos, 1993), and
introduction or reintroduction of Artemia at the outset of salina operation or
after disturbances have decimated the original population. Native Artemia
populations unable to remove significant quantities of particulates, microor-
ganisms, and organic matter can be replaced with more aggressive strains
of the animals (Tackaert and Sorgeloos, 1993). 3 Numbers of brine shrimp
greater than required for efficient production of high quality salt at design
capacity can be harvested (Tackaert and Sorgeloos, 1993).
In crystallizers, brine depths near the lower extreme of the 30-50 cm range
have improved crystal size and increased salt floor integrity and saltfield
efficiency (Butts, 1977; McArthur, 1979; original data). These effects may be
due to increased oxidation of organic matter by aerobic halobacteria at the
shallower depths (Davis, 1990).
Deleterious effects of accumulated organic matter in crystallizers of sea-
sonally operated saltworks can be controlled by flushing mucilage and silt
from the ponds with seawater after harvest, and in crystallizers of continuous-
ly operated salinas by periodic removal and replacement of salt floors. Effects
of petroleum products can be reduced by fast removal of leaking equipment
and immediate cleanup of spilled fuel, grease, or oil.
345

Notes

1Most managers of solar saltfields use specific gravity (S.G.) as an indirect measure of salinity,
and it is for this reason the S.G. is used in this paper rather than salinity per se.
2These values may be lower in some saltfields.
3If this is to be done, however, consideration should be given to conserve natural Artemia
populations (see Geddes and Williams, 1987).

References

Baha AI-Deen, B. and Baha A1-Deen, A. 1972. Posible effecto de microalgas en la forma de
cristalizacion del cloruro de sodio en la salina de Araya. Boletin del Instituto Oceanografico
Universidad de Oriente 11: 35-38.
Barcelona, M.J. and Atwood, D.K. 1978. Gypsum-organic interactions in natural seawater:
effect of organics on precipitation kinetics and crystal morphology. Marine Chemistry 6:
99-115.
Berland, B., Le Campion, T. and Campos Baeta Neves, M.H. 1989. Interaction de la salinit6
et de la temptrature sur la morphologie, la croissance et la composition cellulaire d'une
Cyanobact&ie halotoltrante (Aphanothece sp.). Botanica Marina 32: 317-329.
Binford, M.W., Deevey, E.S. and Crisman, T.L. 1983. Paleolimnology: an historical perspective
on lacustrine ecosystems. Annual Review of Ecology and Systematics 14: 255-286.
Bremer, J.I. 1983. Solar salt production at Exportadora de Sal. Sixth International Symposium
on Salt, Vol. 2, pp. 401-406.
Britten, R.H. and Johnson, A.R. 1987. An ecological account of a Mediterranean salina: the
Salin de Giraud, Camargue (S. France). Biological Conservation 42: 185-230.
Brock, T.D. 1976. Halophilic-blue-green algae. Archiv ftir Microbiologie 107:109-111.
Burnard, E. and Tyler, J.P. 1993. Brine quality management in solar salt operations. Seventh
Symposium on Salt, Vol. 1, pp. 503-508.
Butts, D.S. 1977. Solar evaporation chemistry of Great Salt Lake brines. In: D.C. Greer (Ed)
Desertic Terminal Lakes. Proceedings of the International Conference on Desertic Terminal
Lakes, Weber State College, Utah Water Research Laboratory, Utah State University,
Logan, pp. 125-129.
Chardard, R. 1990. Nouvelles observations sur la structure et la composition du cell-coat de
Dunaliella bioculata (algue verte). Cryptogamie Algologie 11:137-152.
Coleman, M.U. and White, M.A. 1993. The role of biological disturbances in the production
of solar salt. Seventh Symposium on Salt, Vol. 1, pp. 623-631.
Davis, J.S. 1978. Biological communities in a nutrient enriched salina. Aquatic Botany 4:
23-42.
Davis, J.S. 1980. Experiences withArtemia in solar saltworks. In: G. Persoone, P. Sorgeloos, Q.
Roels and E. Jaspers (Eds) The Brine Shrimp Artemia, Vol. 3. Ecology, Use in Aquaculture.
Universa Press, Wetteren, Belgium, pp. 51-55.
Davis, J.S. 1990. Biological management for the production of salt from seawater. In: J.
Akatsuka (Ed) Introduction to Applied Phycology. SPB Academic Publishing, The Hague,
pp. 479-488.
Davis, J.S. 1993. Biological management for problem solving and biological concepts for a
new generation of solar saltworks. Seventh Symposium on Salt, Vol. 1, pp. 611-616.
DeMedeiros Rocha, R. and Camara, M.R. 1993. Prediction, monitoring and management of
detrimental algal blooms on solar salt production in north-east Brazil. Seventh Symposium
on Salt, Vol. 1, pp. 657-660.
Dubessy, J., Geisler, D., Kosztolanyi, C. and Vernet, M. 1983. The determination of sulphate
in fluid inclusions using the M.O.L.E. Raman microprobe. Application to a Keuper halite
and geochemical consequences. Geochimica et Cosmochimica 47: 1-10.
346

Geddes, M.C. and Williams, W.D. 1987. Comments on Artemia introductions and the need for
conservation. In: P. Sorgeloos, D.A. Bengtson, W. Decleir and E. Jaspers (Eds) Artemia
Research And Its Applications, Vol. 3, Universa Press, Wetteren Belgium, pp. 19-26.
Geisler, D. 1981. Gen~se et 6volution des gypses des marais salants de Salin-de-Giraud (Camar-
gue). Bulletin de Minrralogie 104: 625-629.
Geisler, D. 1982. De la met au sel: les facies superficials des marais salants de Salin-de-Giraud
(Sud de la France). Grologie mrditerranrene 9: 521-549.
Gibor, A. 1956. Some ecological relations between phyto- and zooplankton. Biological Bulletin
111: 230-234.
Ginsburg, M., Weizinger, G., Cohen, M. and Ginzburg, B. 1990. The adaptation of Dunaliella
to widely-differing salt concentrations. Journal of Experimental Botany 41: 685-692.
Giordano, M. 1992. Effetti di differenti condizioni di coltura sulla biologia della microalga
verde alotollerante Dunalietla satina. PhD thesis, University of Genoa, Italy.
Giordano, M., Davis, J.S. and Bowes, G. 1994. Organic carbon release by Dunaliella salina
(Chlorophyta) under different growth conditions of CO2, nitrogen, and salinity. Journal of
Phycology 30: 249-257.
Golubic, S. 1980. Halophily and halotolerance in cyanophytes. Origin of Life 10: 169-183.
Helyar, K.R. and Brown, A.L. 1976. Octan-I-ol extraction of molybdophosphoric acid in the
colorimetric determination of orthophosphate. Soil Science Society of America Journal
40: 43-46.
Huntsman, S.A. 1972. Organic excretion by DunalieUa tertiolecta. Journal of Phycology 8:
56-63.
Javor, B. 1983. Planktonic standing crop and nutrients in a saltern ecosystem. Limnology and
Oceanography 28: 153-159.
Jones, A.G., Ewing, C.M. and Melvin, M.V. 1981. Biotechnology for saltfields. Hydrobiologia
82: 391-406.
Jones, J.H. and Yopp, J.H. 1979. Cell wall constituents ofAphanothece haIophytica. Journal
of Phycology 15: 62-66.
Landry, J.C. and Jaccard, J. 1982. Chemie des eaux libres dans le marais salant de Salin-de-
Giraud (Sud de la France). G6ologie M6diterranrene 9: 329-348.
Loeblich, L.A. 1972. Studies on the brine flagellate Dunalielta salina. PhD thesis, University
of California, San Diego.
McArthur, J.N. 1979. An approach to process and quality control relevant to solar salt field
operations in the northwest of Western Australia. Fifth Symposium on Salt, Vol. 2, pp. 325-
334.
Mohr, V. and Larsen, H. 1963. On the structural transformations and lysis of Halobacterium
salinarium in hypotonic and isotonic solutions. Journal of General Microbiology 31: 267-
280.
Nixon, S. 1970. Characteristics of some hypersaline ecosystems. Ph.D. thesis, University of
North Carolina, Chapel Hill.
Oren, A. and Dubinsky, Z. 1994. On the red coloration of saltern crystallizer ponds. II.
Additional evidence for the coloration of halobacterial pigments. International Journal of
Salt Lake Research 3: 9-13.
Rahaman, A.A., Ambikadevi, M. and Sosamma-Esso. 1993a. Biological management of Indian
solar saltworks. Seventh Symposium on Salt, Vol. 1, pp. 633-643.
Rahaman, A.A., Sosamma-Esso and Ambikadevi, M. 1993b. Hydrobiology of two solar salt-
works in India-II. Seventh Symposium on Salt, Vol. 1, pp. 645-655.
Sammy, N. 1983. Biological systems in north-western Australian solar saltfields. Sixth Sym-
posium on Salt, Vol. 1, pp. 207-215.
Sarig, S., Glasner, A. and Epstein, J.A. 1975. Crystal habit modifiers. I. The relationship
between the structure of the additive and the crystal lattice. Journal of Crystal Growth 28:
295-299.
Sarig, S. and Tartakovsky, F. 1975. Crystal habit modifiers. II. The effect of supersaturation on
dendritic growth. Journal of Crystal Growth 28: 300-305.
347

Tackaert, W. and Sorgeloos, E 1993. The use of brine shrimpArtemiain biological management
of solar saltworks. Seventh Symposium on Salt, Vol. 1, pp. 617-622.
Takebe, T., Incharoensakdi, A., Arakawa, K. and Yokota, S. 1988. CO2 fixation rate and rubisco
content increase in the halotolerant Cyanobacterium Aphanothece halophytica grown in
high salinities. Plant Physiology 88:1120-1124.
Tindall, D.R., Yopp, J.H., Miller, D.M. and Schmid, W.E. 1978. Physico-cbemical parameters
governing the growth ofAphanothece halophytica (Chroococcales) in hypersaline media.
Phycologia 17: 179-185.
Van Rijn, J. and Cohen, Y. 1983. Ecophysiology of the cyanobacterium Dactylococcopsis
salina" effect of light intensity, sulphide and temperature. Journal of General Microbiology
129: 1849-1856.
Walsby, A.E., Van Rijn, J. and Cohen, Y. 1983. The biology of a new gas-vacuolate cyanobac-
terium, Dactylococcopsis salina sp. nov., in Solar Lake. Proceedings of the Royal Society,
Series B217: 417--447.
Ward, D.M. and Brock, T.D. 1978. Hydrocarbon biodegradation in hypersaline environments.
Applied and Environmental Microbiology 35: 353-359.

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