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ISSN: 2008-8019

Int. J. Aqu. Sci; 1(1):19-27, 2010

-Review-

The Brine Shrimp Artemia and hypersaline environments microalgal


composition: a mutual interaction
Fereidun Mohebbi
Iranian Artemia Research Center, Urmia, Iran

Abstract
Hypersaline environments are essential, integral and dynamic part of the biosphere. Their management and
protection depend on an understanding of the influence of salinity on biological productivity and community
structure. The aim of this study was to review the relationships between the two basic biological components
of hypersaline environments (micro-algae and Artemia) to provide a better understanding the dynamics of
these unique ecosystems. Algal composition as the main food source of Artemia determines Artemia growth,
reproduction rates, brood size, density, lipid index and cysts yields. Furthermore, the reproduction mode of
Artemia depends on food levels, so that at the low food levels the main reproduction going into cysts. On the
other hand, seasonal fluctu-ations of algal abundance influence Artemia population in temperate large
hypersaline lakes such as the Great Salt Lake (USA), Urmia Lake (Iran) and Mono Lake (USA). However,
Artemia grazing pressure has significant effects on microalgal density.
Key Words: Artemia, hypersaline, microalgal composition, Dunaliella

Introduction such environments are affected by both human


Hypersaline environments are important natural activities and natural dynamics. For example,
assets of considerable economic, ecological, recent climatic changes have led to increasingly
scientific and natural value. These ecosystems intense monsoon rains in some regions and to
span large areas worldwide, not only in salt decreasing precipitation levels in others (Das
production areas (solar salt works, salterns or Sarma, 2007). However, increased drought in arid
Salinas) but also in natural lakes and lagoons, and and semi-arid regions of the world, which most of
in tidal ponds (Javor, 1989). Hypersaline the inland hypersaline lakes are located on, had
environments of both marine and continental dramatic effects on these sensitive ecosystems. In
origin are essential, integral and dynamic part of fact, inland hypersaline lakes within closed
the biosphere while the biogeochemical processes hydrologic basins are subjected to natural and
occurring in their unique ecosystems have induced fluctuations in size and salt concentration
considerable environmental, social and economic over both short and long term intervals (Herbst
values (Shadrin, 2009). Interestingly, their unique and Blinn, 1998).
physical and chemical characterization and Hypersaline environments are generally defined as
distinctive biota set them apart from other aquatic those containing salt concentrations in excess of
ecosystems (Naceur et al., 2009). Hypersaline sea water - 3.5% total dissolved salts (Das sarma
environments and biodiversity associated with and Arora, 2001). Velasco et al (2006) studied the
( ) mohebbi44@yahoo.com
Mohebbi (2010)

effects of salinity changes on the biotic necessary, the abiotic parameters have been
communities in a Mediterranean hypersaline argued. So, the objective of the present study was
stream. They suggested that the number of taxa, to attempt to provide a review based on the
Margalefʼs index and Shannonʼs diversity index recent and previous studies on relationships
decreased with increasing salinity. Their results between the two basic biological components in
supported the initial hypothesis that dilution hypersaline environments i.e. microalgae and
causes an increase in richness and biotic diversity, Artemia in order to provide a better understanding
but a reduction in abundance. In other words, as the dynamics of these unique ecosystems (Jellison
noted Borowitzka (1981): the more saline the and Melack, 1988).
water, the lower the species diversity and simpler
the structure of the system. The number of Algal composition
studies that has been performed on Artemia- Both prokaryotic and eukaryotic algae grow, and
phytoplankton ecological relationships in natural contribute to primary production in high salt
biotopes is very limited; in most cases these concentration waters (Borowitzka, 1981).
studies refer to small-sized salt lakes, or to man-
made salt ponds (e.g. Herbst, 2006; Tanner et Solar saltworks
al., 1999; Warnock et al., 2002). Persoone and The cyanobacterial mats of the solar lakes can be
Sorgeloos (1980) and Lenz (1987) reviewed divided into two layers, the upper layer was
literature data on the productivity of Artemia dominated by coccoide forms such as
habitats in natural environments, both inland and Aphanothece stagnina (Sprengel) A.Braun, 1865
coastal salt lakes. and Synechococcus sp., while the lower was
Management and protection of these changing dominated by filamentous species composed
ecosystems depend upon an understanding of the mainly of Oscillatoria and Spirulina species
influence of salinity on biological productivity and (Gamila, 1999). On the other hand, the
community structure. In other words, salinity microalgae of solar saltworks may be classified
changes both directly or indirectly affects the according to salinity tolerance, namely halobiont
primary production which in turn may influence (salinity range 115 to 185 ppt) such as Anacystis,
Artemia, as the major macrozooplankton of Dunaliella and Spirulina; halophilic (which can
hypersaline waters. On the other hand, Artemia tolerate salinity range of 41 to 150 ppt) including
population is able to influence phytoplankton Lyngbya, Oscillatoria, Gloeocapsa, Synura,
composition by feeding on them; therefore there Amphora, Navicula spp. Nitzschia spp. and
is a continuous reciprocal interaction between Surirella; stenohaline which are abundant below
Artemia and phytoplankton population in 41 ppt, e.g. Chaetoceros, Gyrosigma, Amphora
hypersaline environments. Although, abiotic marina W.Smith, 1857, etc (Rahaman, 2006).
parameters such as temperature, salinity etc. Williams (1981) suggested that the dissolved
which form a background for biological activities oxygen concentrations may well be important in
and governed on these interactions, but since determining salinity tolerance in halobionts and in
these ecological factors have been studied more certain halophiles. Dolapsakis et al (2005)
frequently, so in this study we only focused on the concluded that in higher salinities Dunaliella salina
interactions between Artemia and algae in (Dunal) Teodoresco, 1904, was able to increase
hypersaline environments. However, when its ratio to more than 50% and displace other

Int. J. Aqu. Sci; 1(1):19-27, 2010 20


The Brine Shrimp Artemia and hypersaline environments…

hypersaline microalgae, possibly as a better though valuable, are tougher for the Artemia to
resource competitor. digest.
Temperate lakes The primary producers of benthic habitat in
In temperate hypersaline lakes, salinity as the hypersaline lakes are dominated by cyanobacteria
major dominant parameter determines phyto- and some diatoms with the occasional green
plankton species composition and diversity. In algae. The benthic primary producers are parti-
most of these lakes, Dunaliella spp. mostly cularly active when the phytoplankton and brine
dominate other microalgae, due to their higher shrimp density decline in the water column and
salinity tolerance. For example, in the Great Salt resulting clear water allows light to penetrate to
Lake it has been found that when salinity was the bottom.
near 200 g/l, the phytoplankton community was Microalgal composition of Urmia Lake is roughly
nearly a monoculture of Dunaliella viridis E.C.- similar to the phytoplankton in the Great Salt
Teodoresco, 1905, and nitrogen fixing Lake, which consists predominantly of Dunaliella,
cyanobacteria were absent or rare in the pelagic with an important fraction of diatoms like
region of the lake (Rushforth and Felix, 1982), but Nitzschia and Navicula (Sorgeloos, 1997).
when salinity decreased to about 50 g/l not only However, other algae such as cyanobacteria have
the density of phytoplankton increased, but also contributed in microalgal composition of Urmia
nitrogen fixing Nodularia sp. was detected in the Lake at more limited extent. For example,
lake. (Wurtsbaugh and Berry, 1990). Stephens Mohebbi et al (2006) reported 3 cyanobacteria
and Gillespie (1976) demonstrated that when the (Anabaena, Oscillatoria and Synechoccus) 2 green
lake salinity was higher and nitrogen-fixing cyano- algae (Dunaliella and Ankistrodesmus) 11 diatoms
bacteria absent, nitrogen deficiency may have (Navicula, Nitzschia, Cyclotella, Symbella,
contributed to the summer decline in Synedra, Pinnullaria, Diatoma, Amphiprora,
phytoplankton. Surirella, Cymatopleura and Gyrosigma). Reyahi
Wurtsbaugh and Gliwicz (2001) suggested that et al (1994) observed 6 cyanobacteria (Anabaena,
the pelagic community of phytoplankton in the Anacystis, Chrococcus, Lyngbya, Oscillatoria and
Great Salt Lake during periods that Artemia were Synechoccus) 4 green algae (Ankistrodesmus,
present as juveniles or adults (from Mars to June) Dunaliella, Monostroma and pandorina) 2 diatoms
was dominated by the unicellular green alga (Amphora and Navicula). These variations may be
Dunaliella viridis E.C.Teodoresco, 1905, with more related to limited and irregular sampling or
than 99% of the numbers of phytoplankton and increased salinity of the lake during recent years
between 84 and 99% of the biovolume. The that likely has eliminated some non-tolerant
densities of this alga were between 2800 and species (Eimanifar and Mohebbi, 2007).
-1
4130 cells ml . Other algae such as diatoms Quantitative analysis of chlorophyll a and algal
Amphiprora sp. And Amphora sp. contributed density indicated that primary production in Urmia
modestly to the biovolume on some dates. In the Lake is lower than that of its sister the Great Salt
Great Salt Lake two genera of cyanobacteria Lake and that Dunaliella is the dominant
namely Aphanothece and Coccochloris with the phytoplankton (more than 95% of the total
Dunaliella spp. along with the dozen or so phytoplankton in number) of both lakes (Van
bacterial species provide the food for grazing brine Stappen et al,. 2001; Gliwicz et al., 1995 and
shrimp. Due to their thick silica cell walls, diatoms Mohebbi et al., 2006).

Int. J. Aqu. Sci; 1(1):19-27, 2010 21


Mohebbi (2010)

Halotolerant algae of the genus Dunaliella are the the mouth, broken into smaller pieces or
most ubiquitous eukaryotic microorganisms in discarded (Borradale and Potts, 1967). Digestion
hypersaline environments which can survive even begins with the mandibles reducing the size of
in saturated salt solutions. In Urmia Lake as the food particles and with the addition of a sticky
second largest hypersaline lake in the world secretion from glands near the mouth. The
Dunaliella was the dominant phytoplankton all secretion helps hold the bolus together while
seasons (Mohebbi et al., 2009). The much more secretions all along the digestive tract add
salinity near saturation (>330 ppt) induced Urmia digestive enzymes (Brown 1960). The food is
Lake Dunaliella to accumulate large amounts of passed into the mouth, through esophagus and
beta-carotene in their chloroplasts (Mohebbi et al., into the combined stomach and intestine.
unpublished data). This is a mechanism by which Nutrients are taken up all along the digestive tract
this alga is able to tolerate stress conditions such before the waste is passed on through the anus.
as high irradiance and salinity (López-Uc et al., (Borradale and Potts, 1967).
2009; Garcia et al., 2007). It seems that algal composition as the main food
More recently, a picoplanktonic (2-3µ) green alga, source of Artemia in both natural habitat and
Picocystis salinarum Lewin, 2000, was described culture media has significant effect on Artemia
from Mono Lake which dominates the growth and reproduction rates. For example
phytoplankton community of the lake (Bruce et DʼAgostino (1980) noted that growth of Artemia
al., 2008). was influenced by both the species of
phytoplankton in the diet and the culture media
Artemia and phytoplankton interactions used to grow the phytoplankton. According to
Brine shrimp in the genus Artemia are the Wurtsbaugh and Gliwicz (2001), in the Great Salt
dominant macrozooplankton present in many Lake both the quality and quantity of the food
hypersaline environments (Wurtsbaugh and source, combined with temperature likely
Gliwicz, 2001). This crustacean often dominates influence growth rate and time to maturation of
food web dynamics in hypersaline environments Artemia, as it does in other ecothermic animals
and its grazing activities control water clarity (e.g. Wurtsbaugh and Cech, 1983). Savage and
(Lenz, 1987; Wurtsbaugh, 1992) and Knott (1998) studied the effects of limnological
consequently they are often introduced into salt factors on parthenogenetic Artemia populations
production facilities to minimize algal blooms from Lake Hayward, western Australia. They
(Sorgeloos et al., 1986). The role of microalgae suggested that the major mechanism controlling
and Artemia in the production of high quality salt nauplius survival and recruitment of Artemia in
have been well documented (Davis, 1980). Lake Hayward was food quality and quantity.
Brine shrimp are non-selective filter feeders Low spring temperatures slow growth rates of
feeding on detritus scraped up from the bottom of Artemia in the Great Salt Lake (Wirick, 1972) and
the water column or on unicellular algae and other Mono Lake (Dana et al., 1995). In the summer,
plankton higher up in the water column. Among when perhaps lowered nutrient inputs and high
other functions the rhythmic beating of the limbs grazing rates (Wurtsbaugh, 1992) decrease
pumps food and water through the median gully phytoplankton abundance, Artemia growth is slow
where the food is strained out by the limbs and (Dana et al., 1995). This occurs despite the fact
retained in the food groove and later moved up to that potential growth rates of well-fed animals are

Int. J. Aqu. Sci; 1(1):19-27, 2010 22


The Brine Shrimp Artemia and hypersaline environments…

maximal at midsummer lake temperatures of 25- (Berthelemy- Okazaki and Hedgecock, 1987), but
28 º C (Vanhaecke and Sorgeloos, 1980). Low food level appears to play a more important role.
brood sizes and lipid indices have also been On the other hand, primary production as food
related to the food limitation of the Artemia in the availability or food levels influences adult Artemia
summer. densities in various hypersaline lakes. The lower
In large temperate lakes such as the Great Salt production in the Great Salt Lake supports peak
Lake, USA, Urmia Lake, Iran and Mono Lake, USA, summer adult densities of Artemia near 3 l-1;
where salinity differences are small and have little whears the more productive Mono Lake has
impact on the ecosystem and seasonal differences densities of 6-8 l-1 in the pelagic region (Conte et
mainly imply temperature variations, generally al., 1988).
Artemia has two generations per year. In fact, High salinity of Urmia Lake during recent years up
Artemia population is declining by lack of to saturation levels (about 350 ppt) has
recruitment, increased predation and thermal diminished Aremia population to less than 1
death (Lenz, 1987). Therefore, large temperate individual m-3 compared to 1 individual l-1 during
lakes indicate explicit seasonal cycles with a high the high water levels (Mohammadi et al., 2009).
predictability in which fluctuations from one year It seems that increased salinity inhibits cyst
to another are limited. The spring burst of egg hatching, limits availability of nutrients especially
(nauplii) production is well suited to saturate the unicellular algae and causes high mortality due to
environment with juveniles that can exploit the excessive stressful condition. Additionally, cyst
abundant phytoplankton food resource present clutch sizes of Mono Lake Artemia are nearly
then. As phytoplankton abundance decreased, double those observed in the Great Salt Lake
nauplii survival declined greatly and the (Dana et al., 1990) and cyst weights are equal to
proportion of cysts produced increased or greater than those in the Great Salt Lake
significantly (Wurtsbaugh and Gliwicz, 2001). In (R.Jellison, Pers.comm).
the Great Salt Lake, early spring, when the Cysts yields in various hypersaline lakes and in
primary producers have recovered from the different years can be influenced by primary
previous yearʼs grazing pressure, is the only time production. For instance, in Mono Lake, Artemia
of high food levels. Therefore, this is the time of produced 2.4 × 106 to 5.1 × 106 cysts m-2 during
mass hatching from diaposing cysts and of intense different years, considerably more than the 6.5 ×
reproduction in Artemia (Wurtsbaugh and Gliwicz, 105 m-2 produced in the Great Salt Lake in 1995
2001). (Dana et al., 1990). This variation is likely
Under low-food conditions, female Artemia have because of a higher rate of primary production in
likely evolved mechanisms to produce cysts. Mono Lake (269-641 g C m-2 yr-1; Jellison and
According to Gliwicz et al., (unpublished data) in Melack, 1993) than in the Great Salt Lake (145 g
controlled laboratory experiments, Artemia C m-2 yr-1; Stephens and Gillespie, 1976).
produced 70-90% cysts when food levels were Decreased salinity of the Great Salt Lake
less than 2.5 µg chl a l-1 and 60-70% nauplii when dramatically declined the Artemia abundance due
-1
chl a levels were > 12 µgl . Other factors such as to the appearance of new predator taxa in
temperature, salinity, photoperiod and brood zooplankton assemblage (Wurtsbaugh and Berry,
number also contribute the mode of reproduction 1990). According to this study, small population of
Artemia and other zooplankton were not able to

Int. J. Aqu. Sci; 1(1):19-27, 2010 23


Mohebbi (2010)

clear the water column, thus chlorophyll a levels mode of reproduction switched from cyst
-1
remained high (5-44 µ g l ) and secchi depths production to hatching eggs at low food levels in
low (<2.2) throughout the year. the lake. On the other hand, Gliwicz et al.,
-1
A single Artemia filters 240 mld and therefore, at (unpublished data) suggested that the Artemia
the average population density of 4 sub-adult and body weight in the Great Salt Lake was
adult individuals per liter, this branchiopod is considerably smaller than that of the Farmington
capable of filtering the entire lake volume once a Bay of the Great Salt Lake, where chlorophyll was
day (Reeve, 1963). Thus the Dunaliella population much higher and Artemia less abundant due to
density remains extremely low, and does the lower salinity.
density of other green algae, diatoms and Artemia raised in laboratory in lake water without
cyanobacteria that are able to reproduce fast added food withheld eggs in egg sacks (Gliwicz et
enough to compensate for grazing losses. al., 1995). Female Artemia are prevented from
According to Gliwicz (2003), in contrast to allocating sufficient resources to reproduction
Dunaliella which is a typical euplanktonic species, below a threshold food concentration, which
many other taxa are not suspended in the lake results in a long inter-brood interval as the
water, but live in refuges where grazing losses are clutches of eggs are withheld in the brood sacs. At
lower. These refuges are provided by the interiors the low food levels only 1 of 13 broods produced
of the long tubular setae of Artemia exoskeleton, was ovoviviparious with the bulk of the
which form the combs on the filtration reproduction going into cysts, and brood size were
appendages. The exoskeleton is shed at each of relatively small (15.6) (mean ± 7.3 ISD) eggs
the 13 or 14 molts necessary for Artemia to attain female-1 day-1; in contrast, in the high food levels
maturity and large quantities float in the water. It 85% of broods were ovoviviparous and mean
was found that diverse algal-cyanobacteria brood size was 53.6 (mean ± 24.8 ISD) eggs
community that colonize long filtration female-1 day-1 (Gliwicz et al., unpublished data).
appendages of Artemia, represented up to 20% of Most of the first cohort Artemia was already adult
the available food for adult Artemia, when the by early May, but at the early June, the density of
preferred free-swimming Dunaliella was at an juveniles was dropped to one tenth of the
extremely low density in the entire the Great Salt expected value. Therefore, 90% of the second
Lake southern arm. Artemia juveniles and nauplii generation hatching from ovoviviparous eggs had
can not readily access the algae colonizing died, evidently because food levels had declined
discarded exoskeleton which induces higher from 25 µg chl a l-1 to less than 1 µg chl a l-1 in
survival in the older instars of Artemia than early June (Gliwicz et al., unpublished data).
younger ones. This is evident from the seasonal
changes in size distribution indicated by the Acknowledgements
densities of discrete size classes (Gliwicz et al., I would like to appreciate Iranian Fisheries
unpublished data). Research Organization (IFRO). I also appreciate
As the food availability declines from June to Professor Wayne Wurtsbaugh for providing
November: the individual Artemia lipid index valuable references.
gradually decreased (Wurtsbaugh an Gliwicz,
2001); juvenile Artemia survival was held much References
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