In: Impact, Monitoring and Management ISBN 978-1-60876-487-7

Editors : Ahmed El Nemr, pp. 2010 Nova Science Publishers, Inc.

Chapter 4

FRESHWATER CYANOBACTERIAL (BLUE-GREEN

ALGAE) BLOOMS: CAUSES, CONSEQUENCES AND
CONTROL

NK Sharma1, KK Choudhary 2, Rakhi Bajpai 2 and AK Rai1 2, *

1
Department of Botany, Postgraduate College,
Ghazipur (UP) -233001, India
2
Department of Botany, Banaras Hindu University,
Varanasi (UP) -221005, India

ABSTRACT
Harmful algal blooms (HABs) are a natural phenomena however, due to human
activities and interventions, incidences of HABs have increased globally. Onset,
development and proliferation of blooms are closely associated with the nutrient
enrichment of water bodies (eutrophication) and climatic changes; and their possible
interaction. Cyanobacteria are amongst the most successful bloom forming algae. They
can convert and use different forms of C, N, P, and S that help them in occupying almost
all kinds of aquatic habitats. Moreover, they grow well in shaded light, show resistance
against grazing pressure and release allelochemicals to out-compete co-occurring
organisms. Presence of gas-vacuoles facilitates their migration in the water column to
ensure enough light and nutrient availability. Cyanobacterial blooms adversely affect
water quality, structure and composition of biological communities and a range of
ecological services. Many of the bloom forming cyanobacteria produce toxins
responsible for mass mortality of aquatic and exposed vertebrate populations. This
chapter describes the causes and consequences of cyanobacterial blooms and a few
measures adopted to control bloom formation and proliferation. The impact of climatic
change on cyanobacterial bloom formation has also been discussed.

1 E-mail: akrai.bhu@gmail.com.
74 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

INTRODUCTION
Phytoplankton production is important in supporting the productivity of aquatic
ecosystems. Biomass produced by the phytoplankton provides the energy and materials for
the use of other organisms in aquatic food webs. The size of phytoplankton assemblages
depends upon the balance between bottom-up and top-down controls. Bottom-up control
operates through level of light and nutrients, which determines the rate of biomass production
of a particular water body. However, grazers consume this newly produced biomass and exert
top-down control [1]. Due to rapid industrialization and other socio-economic factors
various types of organic and inorganic pollutants are released into water bodies bringing a
change in their nutrient status, pH, turbidity and temperature. Consequently, accelerating the
bottom-up effect weakens the top-down control, resulting in massive phytoplankton growth
i.e., the bloom.
In late June 2008, the waters and shores at the Qingdao venue hosting the Olympic
sailing regatta experienced a massive green tide covering about 600 sq km. Lasting over two
weeks, it took more than 10,000 people to clean up, removing over one million tonnes of
green mass from the beach and coast [2]. Organisms mainly responsible for such changes are
algae. For instance, in the above case it was the green alga Enteromorpha prolifera. Luxuriant
growth of algae makes water colored and turbid. A condition often termed as blooming of
water bodies. A bloomed water body (chlorophyll-a 300g/L) has algal biomass higher than
that of oligotrophic (1.5 to 10.5g/L) waters [3]. It has a minimum cell concentration of
approximately 20,000 cells mL-1 [4].
Blooms are not static communities, rather they show variation in space and time. They
exhibit a succession of dominance; diatoms and green algae dominate in the winter and
spring; green algae dominate in late spring and summer, and cyanobacteria dominate in late
summer fall. There are a number of taxonomically unrelated algal species that form blooms,
which are identified by the dominant group/species e.g. cyanobacterial bloom or Microcystis
bloom [4]. Eutrophic (nutrient rich, highly productive system) and hypertrophied (extreme
eutrophy) conditions favor as well as extend the period of cyanobacterial (blue-green algae)
dominance in an ecosystem [3]. Cyanobacteria are the most notorious bloom formers [1].
They infest almost every type of waters (with the possible exception of acidic waters) but,
freshwater cyanobacterial bloom especially; waters used for drinking and recreational
purposes are of the greatest concern. Since a number of surface bloom forming cyanobacteria
produce toxins, there are hazardous effects on co-occurring aquatic biota as well as exposed
human and animal populations [5].

1. CYANOBACTERIAL BLOOMS
Cyanobacteria constitute a group of prokaryotic photoautotrophs with an oxygenic mode
of photosynthesis. They are supposed to have originated and evolved in the proterozoic era
in nutrient limiting conditions; and subsequently distributed into a wide range of habitats [6].
They show morphological, reproductive and physiological adaptations in order to occupying
different ecological habitats. Cyanobacteria form symbiotic associations with other microbes,
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 75

higher plants and animals and produce different kind of resting spores. Some genera are able
to fix dinitrogen (N2) in adverse nitrogen-limiting condition.
Cyanobacteria possess certain unique adaptations that make them a successful competitor
in a bloom environment. These include, their ability to grow in warm waters, capture reduced
photosynthetic flux densities, utilize low TN: TP ratio and, to access low dissolved CO2
concentration (in form of bicarbonate). In addition, bloom forming cyanobacteria may
undergo organizational changes during different stages of bloom development. For example,
filamentous forms such as straight, spiral or twisted chains of cells may achieve secondary
structures due to aggregation or enlargement of filaments [7, 8]. Similarly, globular colonies
change to loose aggregation of separate colonies [9]. Ability to reorganize thallus structure
according to environmental conditions helps cyanobacteria in buoyancy regulation, which is
an active process in which gas vacuoles inflate and deflate in an effort to regulate the cells at
an optimum depth for nutrient and light availability. Vertical migration also helps in
formation of blooms at the surfaces. Cyanobacteria commonly inhabiting eutrophic
freshwater bodies are species of Microcystis, Oscillatoria, Spirulina, Anabaena,
Anabaenopsis, Aphanizomenon, Nodularia, Cylindrospermum, Tolypothrix, Calothrix,
Mastigocladus, Scytonema, Nostoc, Rivularia and Gloeotrichia [10].

1.1. Temporal and Spatial Dynamics of Cyanobacterial Blooms

In a bloomed aquatic ecosystem, population size, species composition and cell densities
of cyanobacteria show temporal and spatial variation. Some freshwater bodies may have
seasonal blooms that start in summers and last into autumn. A few have persistent blooms,
while in many cases blooms occur as extreme peaks and crashes lasting a few days or weeks
[11]. Seasonality of blooms in a particular water body is subject to the extent to which
different environmental factors influence bloom dynamics [11]. In temperate regions blooms
generally occur during the warm, windless days of late summer (possibly due to increased
light intensities and temperature). Havens [11] reported that in deep temperate eutrophic lakes
with stable summer stratification, phytoplankton progresses through the dominance of
diatoms in spring, followed by clear water; cyanobacteria dominate in mid to late summer
with water temperature >200C, depletion of inorganic nitrogen and free CO2. In tropics,
cyanobacterial blooms may appear at any time as temperature remains relatively constant
throughout the year. If winters are not too cold, cyanobacterial blooms may occur throughout
the year [12, 13]. In such conditions, diversity and dominance of species are determined by
the thermocline establishment brought about by meteorological changes. Depression in the
thermocline increases diatoms populations and decreases cyanobacterial populations. Re-
establishment of thermocline again makes cyanobacteria abundant. Reynolds [14] found that
succession of algal species in eutrophic water bodies is determined by stratification and
nutrient availability. In general, in eutrophic water bodies following successional sequence
has been observed: diatoms cyanobacteria green algae dinoflagellates, depending
upon light and nutrient availability [15]. Eutrophy favors the growth of diatoms and
cryptomonads but cyanobacteria can maintain their dominance even under reduced nutrients
[15].
Cyanobacterial blooms may develop within (metalimnatic bloom) or onto the surface
(surface bloom) of the water bodies. However, based on perceived threat to human, surface
76 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

blooms are of greater concern compared to metalimnetic blooms [1]. Although appearance of
surface blooms is a seasonal event, persistent surface blooms have now become common in
many eutrophic water bodies. This may be because of the loss of buoyancy regulation and
senescence [16], which supports the cyanobacterial dominance at the surface [17, 18]; or due
to the presence of underlying colonies that often blocks the downward movement of
cyanobacteria [16, 19]. For instance, in eutrophic water bodies metalimnion is dominated by
diatoms and prevent the downward migration of cyanobacteria. Many planktonic
cyanobacteria contain gas vacuoles that help cyanobacteria in buoyancy regulation [20]. A
gas-filled vesicle has a density of about one tenth that of water making the cell lighter than
water [21]. Gas-vacuoles remain under dynamic state (i.e., vacuole formation vacuole
collapse) depending upon environmental changes. Buoyancy regulation enables cells/colonies
to acquire light and nutrients in water column therefore, provides a competitive edge over
other phytoplankton. In aquatic systems, when buoyancy is reduced, small-sized
cyanobacteria dominates while, large-sized are prevalent at enhanced buoyancy. Turbulence
is another factor facilitating the quick appearance of surface bloom due to upward migration
of an existing dispersed phytoplankton communities present in metalimnion [22].
There are some cyanobacteria inhabiting both surfaces as well as metalilimnionic waters.
For example, cyanobacteria Aphanizomemon flos-aquae and Oscillatoria agardhii form
surface bloom but, are also reported from metalimnion of eutrophic lake [23]. Both these
species occur as single filaments in metalimnion but as aggregates in an epilimnetic zone
[23]. Likewise, within a genus, species may show variation in their spatial distribution. For
example, Oscillatoria rubescens occurs in the metalimnetic zone while, O. agardhii may
occur in metalimnetic zone [24, 25, 26] as well as in mixed layers [27]. In general,
cyanobacterial population gradually decreases with water depth but, diatom population
increases with increase in water depth. This contrasting depth distribution helps
cyanobacterial population to remain in epilimnion.

1.2. Factors Affecting Cyanobacterial Bloom Formation and Proliferation

The exact mechanism behind the development of cyanobacterial blooms is still to be fully
resolved. Factors such as environmental (nutrient enrichment, light intensity and duration,
temperature, alkalinity), biotic (zooplankton grazing, viral lysis, allelopathic interactions) and
water body characteristics (physical structure, residence time, water turbulence) are supposed
to be involved in the development and expansion of freshwater algal blooms [4]. Paerl [10]
opined that physical factors are mainly responsible for the development of cyanobacterial
bloom in marine ecosystems, while trophic status is more important for freshwater
ecosystems. Based on trophic state (nutrient status and productivity), natural aquatic bodies
are categorized as oligotrophic, mesotrophic and eutrophic. Oligotrophy symbolizes a nutrient
poor state of the water body while; mesotrophy is nutrients rich state providing optimal
conditions for growth and development of species. Eutrophy (excessive nutrients) is a
temporary state released with the utilization of nutrients by phytoplankton, and leads to
increased productivity of the water body. Due to increased surface: volume ratio, algae utilize
nutrients more efficiently than that of other phytoplankton groups leading to exuberant
population growth in nutrient enriched conditions [7]. However, behavior of different
cyanobacteria in natural ecosystems is not similar, and is dependent upon their
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 77

ecophysiological properties. Cyanobacteria with similar eco-physiological characteristics

(ecostrategists) are grouped together; different ecostrategists occupy different ecosystems as
well as different niches of an aquatic ecosystem. Knowledge about the factors (Table 1)
responsible for cyanobacterial bloom development is important in developing control
measures.

Table 1. Physico-chemical conditions favoring surface cyanobacterial bloom formation

Factors Conditions
Nutrient input Excessive P loading
Water temperature High (>25OC) temperature
Water column Persistent and stable water column; depends upon size and volume of
the system and wind velocity
Flushing Long water retention time/ slow flushing
Dissolved carbon Low dissolve inorganic carbon support cyanobacteria growth
N:P ratio Low (<20)
Light High irradiance and long day length
Salinity Low
pH High pH (>8)
Large-scale vertical mixing prevents surface accumulation of blooms,
Vertical mixing favors eukaryotic taxa
Shears (small-scale turbulence) inhibit cyanobacterial bloom
formation

1.2.1. Nutrient Enrichment

Eutrophication is a natural aging process of aquatic ecosystems resulting from hundreds
or thousands of years of human activities that added nutrients to them [28]. Nixon [29]
defined eutrophication as the process of increased organic enrichment of an ecosystem
generally through increased nutrients inputs. In general, eutrophication favors cyanobacterial
bloom formation however, after a certain point eukaryotic algae replace the cyanobacteria
[11, 30]. Concentration of individual nutrients, their modifications and relative ratios have
long been held responsible for the development of cyanobacterial blooms [31]. Amongst
various nutrients, nitrogen (N) and phosphorous (P) have mainly been implicated in
cyanobacterial blooming of water bodies. Other nutrients include potassium, sodium, and
magnesium salts; depending upon the requirement of individual species/groups. Provasoli
[32] considered monovalent ions as absolutely essential for algal growth and development.
Industrialization as well as intensive agricultural practices has led to the substantial increase
in the concentration of such nutrients in waters (i.e., allochthonous sources, Table 2). Besides
these outside sources, nutrient content of an ecosystem can also increase due to internal
recycling of nutrients (i.e., autochthonous source). Rate of water renewal plays a critical role
in nutrient loading and subsequent eutrophication. Stagnant waters are more sensitive to
eutrophication than running waters, since such waters collect excessive nutrients from other
sources than replenished waters. Multiple cloudy days and intense rainfall which increases
flushing rate reduces the bloom development [11]. The drying of wetlands also increases
nutrient concentration leading to eutrophication [33]. Photosynthetic depletion of free CO2
from waters favors cyanobacterial dominance in blooms. Since cyanobacteria have low Ks for
78 NK Sharma, KK Choudhary, Rakhi Bajpai et al.
CO2 and can use bicarbonate as carbon source [34]. It also allows cyanobacteria to form
surface bloom as they move to the air-water interface where CO2 is available in plenty.
Below, we have discussed the importance of nitrogen and phosphorus in cyanobacterial
bloom formation.
Table 2. Allochthanous sources of nutrient enrichment and their importance
Type Characteristics Example
Point sources When nutrients enter Industrial and domestic
into water bodies wastes;
directly from their
source of origin; Wastewater treatment plants
Storm water drains
Non-point Nutrients are gathered Soil Retention
sources at one site and carried (Excess application of
away to the water chemical fertilizer to
bodies by other means agricultural fields often
accumulates in soils and
eventually makes its way
along with rain water to
water bodies causing
eutrophication)
Runoff to surface water and
leaching to groundwater
(Excess nutrients move
either horizontly into other
surface waters i.e., runoff or
vertically downwards to
ground waters i.e.,
percolation or leachate)
Atmospheric deposition
(Due to combustion of fossil
fuels different compounds
are released into the
atmosphere and deposited
onto water bodies e.g.,
nitrogen is released into air
due to volatilization of
ammonia and nitrous oxide
production, which eventually
moves into water bodies
along with rain i.e., acid
rain).
Importance
Point sources are less
important and
contribute a little in
eutrophication
because they may be
controlled and
regulated with little
efforts
They are difficult to
regulate and vary
with season,
precipitation and
other irregular events.
For N and P, non-
point sources act as a
major input in many
water estuaries [29,
109].
Sharpley et al. [110]
reported that in many
water bodies amount
of phosphorus lost to
surface waters
increased linearly
with the amount of
phosphorus in the
soil.
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 79

1.2.1.1. Nitrogen
Cyanobacterial blooms occur mainly in eutrophic waters. Therefore, it was assumed that
they require high phosphorous and nitrogen [35]. Amongst the various nutrients affecting the
growth rate of cyanobacteria induced eutrophication; nitrogen has been the most limiting in
water bodies [36]. In natural ecosystems, nitrogen is available in different inorganic forms
viz. NO3-, NO2--, NH4+ and organic forms like amino acids, organo-nitrates and urea [37, 38].
Concentration and available forms of nitrogen regulate the species composition and relative
size distribution of the phytoplankton cells in communities [39]. In oligotrophic lakes, N2-
fixing cyanobacteria appear first and colonize the system, while in eutrophic water bodies,
species deficient in nitrogen will appear first and utilize nitrogen with enhanced biomass
production [40]. Large-size species will have maximum accumulation leading to their
abundance [41, 42]. Similarly, NH4+ rich waters are generally dominated by small species as
they have greater affinity for NH4+ over NO3- than large species [43].

1.2.1.2. Phosphorus
In freshwater bodies, availability of phosphorous (P) is the most crucial factor
determining the growth of cyanobacteria. Cyanobacteria take P in orthophosphate form (PO4-
3
), and are bestowed with an ability to convert other phosphate (non-usable, non-available)
forms into usable orthophosphate form. High concentration (30-100g L-1) of total
phosphorous promotes cyanobacterial bloom formation [44]. Moreover because of their high
affinity (i.e., growth under P-limited condition) for P, they can store substantial amount of P
during P-sufficient condition to continue for a few generation (cell division). Alternatively,
excess P-loading in the water body may facilitates the growth of other phytoplankton leading
to increased turbidity (low light availability), which favors cyanobacterial growth [35].
Besides alone, phosphorous in combination with other nutrients may regulate
cyanobacterial dominance in bloom environment. Ratio of nitrogen and phosphorous (N:P
ratio) is another measure regulating cyanobacterial bloom formation. Redfield ratio (i.e.,
16N:1P) is a measure of an appropriate N: P ratio for the sustainable development of aquatic
ecosystems. Eutrophic water bodies with low TN: TP ratios favor cyanobacterial dominance
[31, 45]. High N: P ratio is an indicative of phosphorus limitation and vice-versa [46].
However, Reynolds [47] questions the tenability of resource ratio hypothesis and discounts
this as coincidental. According to Downing et al. [44], TP is better predictor of cyanobacterial
dominance than TN: TP ratios. Further, P is limiting nutrients in freshwaters while, N is in
estuarine and marine waters [1, 48].

1.2.2. Environmental Factors

Environmental factors such as light, temperature, water column stability and pH are
important determinants of cyanobacteria bloom development [11, 34]. In aquatic ecosystems,
the zone in which photosynthesis occurs is known as euphotic zone (Zeu). It extends from the
surface to the depth which receives 1% of the surface light intensity. It may be deeper or
shallower than the mixed upper zone of a thermally stratified water body (i.e., the epilemnion,
Zm). Photosynthesis driven high biomass production often results in turbidity of the water.
Consequently, the eutrophic zone becomes more shallow than the epilimnion of the water
body (Zeu/Zm ratio is <1) [35].
80 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

Biliproteins i.e., allophycocyanin, phycocynin and phycoerythrin (absorption range 500-

650 nm) together with chlorophyll-a allow cyanobacteria to grow at low irradiance (turbid
waters) [13]. In general, light intensity at the surfaces of water bodies range from 700 to 1000
Em-2s-1. Light intensity of 320 E m-2 s-1 is lethal for many cyanobacteria. However, if
exposed intermittently, even under high-light intensity cyanobacteria grow to their near
maximal rate [49].Van Liere and Mur [50] reported that exposure (> 180E m-2 s-1) to
extended period inhibited the growth of Oscillatoria agardhii (now Planktothrix). Under low
intensity light conditions cyanobacteria out-grow other phytoplankton species. For instance,
Van Liere and Mur [51] observed that in a similar but low-light condition, cyanobacterium
Oscillatoria agardhii out-competed the green alga Scenedesmus protuberance (requires high
intensity light). Surface-blooms forming cyanobacteria are more tolerant to high light
intensities than their sub-surface dwelling counterparts. Possibly, because of high production
of carotenoids which protects the cells from photoinhibition [52].
Cyanobacteria have low maintenance cost in terms of energy. At low light intensity their
growth rate is higher compared to other phytoplankton species. This provides for them a
competitive edge in turbid water bodies. Kallqvist [53] reported that in a eutrophic lake, at
one meter depth, diatom such as Asterionella, Synedra and Diatoma grew faster than the
cyanobacterium (Planktothrix). However, at intermediate depth (2 m) growth rate was similar
for all the species. While, in deep waters, (3 m) only Planktothrix grew.
High pH favors cyanobacterial bloom formation [11]. Likewise, they attain maximal
growth at temperatures range 25-30C [54]. This temperature is high enough for the growth
of green algae and diatoms: a possible explanation as to why cyanobacterial bloom occurs
during summer in temperate and boreal water bodies. Zhang and Prepas [55] argued that
thermal stratification of the water column is important for the formation of cyanobacterial
blooms. Further, stochastic processes such as wind velocity and rainfall affect water column
stability. Low wind velocity and absence of rainfall facilitates cyanobacterial bloom
formation. The effects of artificial mixing (water column stability) on phytoplankton have
been investigated by several groups. Visser et al. [56] reported that artificial mixing prevents
blooms of Microcystis sp.

1.2.3. Biotic Interactions

In bloom, cyanobacterial populations are relatively stable. As such, they show resistance
against well known zooplankton grazers (except, some ciliates and rhizopods) but are
susceptible to viral, bacterial and actinomycete attack. However, the significance of such
grazing to cyanobacterial population breakdown is not very well understood [35]. Moreover,
buoyancy regulation further lowers the sedimentation of cyanobacterial populations. Hence,
once a population is established, it shows high prevalence [35].
Cyanobacteria produce various kinds of secondary metabolites that may have allelopathic
properties. Allelopathy refers to the inhibitory or stimulatory effect of organisms through the
production and release of diverse kind of organic compounds. This phenomenon is closely
related with the competition for limiting nutrient resources. Allelochemicals are usually
produced under nutrient limited condition [57]. Arguably, under cultural eutrophication,
availability and altered nutrient ratios can stimulate production of allellochemicals [57].
Suikkanen et al. [58] believed that allelochemicals produced by Microcystis aerugenosa may
be responsible for their dominance (suppression of other phytoplankton) in blooms.
Cyanotoxins such as microcystins may affect the structure, composition, succession and
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 81

pattern of species dominance in a phytoplankton community [59, 60]. For instance,

researchers believe that persistent bloom of Cylindrospermopsis raciborskii in Lagoa Santa
Lake (Brazil) could be because of the allaelochenmical produced by the species [61, 62].
Suikkanen et al. [63] found that cell-free extract of Aphanizomenon and Nodularia stimulated
the growth of the cyanobacterial community.

1.2.4. Water Body Characteristics

In shallow lakes (mean depth <2 m), cyanobacterial bloom may persist for years provided
the ratio of euphotic depth to mixed epilimniotic depth (Zeu/Zm) never falls to the level that
inhibits the growth of low-light adapted cyanobacteria [64], but remains low enough to
exclude high-light adapted plankton [11]. Zohary et al. [65] reported that in a deep water
body, high irradiance and low wind velocity result in short-lived scum of Microcystis
aeruginosa that crashed during periods of high flushing and washed away, allowing other
algae to dominate. Growth rate of cyanobacteria is usually much lower than that of other algal
species [51, 66]. Therefore, they require long water retention times to form bloom in the
waters. Water bodies with short retention times usually do not experience cyanobacterial
bloom [35]. Tsujimura and Okubo [67] linked development and reoccurrence of summer
bloom in an eutrophic lake to the emergence of sediment akinete population.

1.3. Strategies Adopted by Cyanobacteria During Bloom Formation

Cyanobacteria vary in their strategy to adopt and occupy all the possible niches present in
a habitat (Table 3). Some cyanobacteria develop large aggregates of cells and filaments
distributed over the water column (i.e., scum-formers). Since, photosynthesis is high at the
water surfaces consequently cells store large quantity of carbohydrates that act as ballast and
induce sinking of the aggregates. Large colonies sink faster than small ones; single cells
remain unaffected. In deep waters, cells utilize carbohydrates during respiration, and also
form new gas-vesicles [68] which assist colonies in returning to the euphotic zone [69].
Under calm conditions of the night, colonies may accumulate on the water surfaces. The
frequency of vertical migration depends upon colony size [35]. In temperate regions, during
autumn photosynthesis overtakes respiration, which allows carbohydrate ballast to persist.
Colonies sink to the bottom, over-winters, and after gradual consumption of carbohydrates
small colonies come up during spring [35]. However, such regulation is only possible in
deeper (> 3m) water bodies.
Blooms of many species remain homogenously dispersed throughout the epilimnion (i.e.,
homogenously dispersed ecostrategists). They are extremely sensitive to high light intensity
and show weak buoyancy regulation. Such types of ecostrategists are found in eutrophic and
hypertrophic shallow lakes forming monoculture but persistent blooms. In addition, a few
cyanobacteria form blooms in metalimnion (i.e., stratifying ecostrategists). The phycoerythrin
pigment of such species allows them to absorb the green light that penetrates up to this depth.
Most metalimnetic blooms are found at light intensities of 1-5% of the surface irradiance
(Zeu/Zm = 0.7-1.2) [35]. The cyanobacterial bloom of ecosystems that are low in inorganic
nitrogen but receive enough light is dominated by N2-fixing species (i.e., N2-fixing
ecostrategists) [10]. Restorative strategies that reduce nitrogen concentration in water bodies
increase the probability of N2-fixing cyanobacteria to dominate in the subsequent bloom.
82 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

Cyanobacteria may also form mats on the bottom sediment of shallow but clear (allow light to
reach to the sediment bottom) water bodies.

Table 3. Cyanobacterial ecostrategists; Mur et al. [35].

Type Habitat Characteristics Genera/species

Scum-formers Calm, buoyancy regulation is
operational
Homogeneously dispersed Vertical mixing less frequent,
shallow, nutrient enriched
Stratified Metalimnion of thermally
stratified lakes
N2-fixers wellstratified eutrophic lakes,
low inorganic nitrogen, but
well lit, intermediate stages of
lakes undergone nutrient
regulation
Small colony formers Small, intermittently flushed
lakes
Benthic Very shallow water bodies
Microcystis, Anabaena,
Aphanizomenon
Planktothrix agardhii,
Limnothrix redekei
Planktothrix rubescens, and
other Planktothrix spp.
Anabaena, Aphanizomenon,
Cylindrospermopsis,
Nodularia, Nostoc spp.
Aphanothece sp.
Oscillatoria limosa

2. CLIMATIC CHANGE AND CYANOBACTERIAL BLOOMS

During the last century, human activities such as fossil fuel burning, changed land use
patterns and deforestation have substantially increased the atmospheric CO2 level. According
to the estimate of an intergovernmental panel on climatic change, the atmospheric CO2 level is
estimated to be more than double by 2100 [70]. Little is known about the way these climatic
changes will affect the formation and proliferation of cyanobacterial blooms. Studies indicate
that global warming and worldwide proliferation of harmful cyanobacterial blooms are
positively linked [71]. Frequency of cultural eutrophication has increased substantially in
recent decades [72]. However, cultural eutrophication is not the only but one amongst
multiple anthropogenic factors responsible for the global expansion of cyanobacterial bloom
[71]. Other factors such as elevated CO2 level and rising temperature may also affect the
development and proliferation of cyanobacterial blooms.
There are a few studies that had investigated the impact of CO2 enrichment on bloom
development [73, 74]. The majority of these deals with the impact of CO2 enrichment on
marine phytoplankton; only a few on freshwater algal blooms [75]. Most water bodies are
supersaturated with CO2 owing to inputs of carbon from deposited sediments (recycling) as
well as fresh input from the catchments [76]. Normally, in natural freshwater bodies,
dissolved inorganic carbon (DIC) concentration remains high enough (1.5-2.2 mmol/L) to
that required for normal cynobacterial growth [77]. However, water bodies experiencing
intense cyanobacterial bloom may suffer from carbon limitation even under high
concentration of DIC [75]. Change in CO2 availability may affect physiology, nutrient cycling
and interactions among major phytoplankton groups [78, 79]. The doubling of the CO2 level
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 83

may increase CO2 dissolution resulting in lowering of the pH values. Concentration and form
of DIC (CO2, H2CO3, HCO3- and CO3 2- ) is strongly linked to the pH value [75]. Efficiency
of RuBisCO differs between different phytoplankton groups [80]. Cyanobacterial RuBisCO
shows low affinity for CO2 (Kc value 200-300 mol/L) [77]. But, cyanobacteria possess
effective CO2-concentrating mechanism (CCM) that elevates CO2 level near the enzyme [81].
Induction of CCMs is regulated by the external concentration of DIC. And, high CO2
concentration of water is likely to repress the CCM.
Cyanobacteria grow better at higher temperatures (>250C) than do diatoms and green
algae [82, 83]. Therefore, rising temperature may provide them competitive advantage over
other planktonic groups [83, 84]. Further, the warming of surface waters strengthens the
vertical stratification and reduces vertical mixing in water bodies [71]. It also lengthens the
optimal growth period (due to global warming lakes stratify earlier in spring and de-stratify
later in autumn). Presence of gas vesicles in many cyanobacteria helps them to exploit
stratified conditions. The dynamic nature of gas vesicles keeps cyanobacteria mobile in the
water column assisting them in getting light and nutrients. They accumulate cyanobacteria at
the surface even when mixing is weak [71, 82, 85, 86]. Also, dense cyanobacterial surface
blooms may even locally increase water temperatures through the intense absorption of light
[87, 88] that could provide additional competitive advantage to cyanobacteria over non-
buoyant phytoplankton [71]. Rising temperature is likely to affect the pattern of precipitation
and drought that could further enhance cyanobacterial dominance [71]. Freshwater discharge
prevents blooms through flushing however, as the discharge recedes and water residence time
increases, nutrient loads will increase, eventually promoting blooms [71].
Excess withdrawal of freshwater for agricultural use is likely to increase the salinity of
freshwater bodies. However, little is known about the impact of increasing salinity on
cyanobacterial bloom development. Marine and brackish cyanobacteria are more salt-tolerant
than freshwater phytoplankton species, [89] as evidenced by increasing reports of toxic
cyanobacterial blooms in brackish waters [90]. Climatic changes have forced some species to
expand their geographical range, and are now reported from previously unknown territories
[86, 91]. Therefore, it is argued that high nutrient loading, rising temperature, enhanced
stratification, increased residence time, and salinazation all favor cyanobacterial dominance in
many aquatic ecosystems [71]. Paerl and Huisman [92] have provided an extensive review of
the effect of climatic chage on global expansion of cyanobacterial blooms.

3. ECOLOGICAL CONSEQUENCES OF CYANOBACTERIAL BLOOMS

Cyanobacterial blooms are responsible for a number of socio-economic as well as
ecological problems (Figure 1). They interfere in the process of sedimentation and often clog
the sand filters used in water treatment plants. Also, they corrode concrete and metallic walls
of the pipes and boilers by secreting acids (carbonic and oxalic acids). Blooms change the pH
and hardness of water by consuming dissolved CO2 hence, rendering it unfit for drinking
purposes. Furthermore, cyanobacterial mucous encourages the growth of pathogenic
organisms such as amoeba and bacteria. Cyanobactrial bloom may affect ecosystems directly
(toxic effect on fish and other aquatic fauna) or indirectly (shading, oxygen depletion etc).
Some of the effects are discussed below.
84 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

Figure 1. A summary of socio-ecological responses and impacts associated with freshwater

cyanobacterial blooms (redrawn from Havens [11]).

3.1. Decreased Water Transparency

Dense surface cyanobacterial blooms causes a decrease in water transparency. In

addition, a regular deposition of particulate matters (nutrient loading) in water bodies together
with metalimniotic cyanobacterial blooms also decreases transparency of the waters (increase
the turbidity). Resultantly, light traveling to the different strata of water bodies filtered out
affecting the growth, and development of aquatic populations. For instance, shading affects
the photosynthesis of the metalimnetic populations responsible for maintaining oxygen in
deep waters. Massive surface blooms also decrease the level of turbulence in water bodies
which, doesnt allow mixing of different strata. Consequently, the oxygen level of
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 85

metalimnion and benthic zone declines. This affects the internal cycling of nutrients and may
cause nutrient deprivation to surface populations.

3.2. Increased Productivity and Decreased Biodiversity of the Water Body

Blooming in a water body increases its productivity. Nutrient enrichment releases the
nutrient-stressed condition prevailing in the oligotrophic water body (P-limitation in case of
cyanobacteria) that facilitates the growth of cyanobacteria. This also triggers the competition
for efficient utilization of nutrient(s) hence, decrease in the biodiversity of the system.
Further, a dense and long-lasting bloom suppresses the growth of submerged epiphytes
through shading (light limitation) [83].

3.3. Decreased Biomass of Benthic Flora

Dense cyanobacterial surface blooms hamper the penetration of light into waters. This
causes death and subsequent settling of cells resulting in the increase of organic carbon
deposition to the bottom of water bodies. Increase organic carbon availability along with
empouring of sewage wastes promotes microbial activity, which create hypoxia (<2 mg O2 L-
1
) or anoxia in bottom-waters [93, 94, 95]. Consequently, bottom waters become unsuitable
for benthic populations. If anoxia persists for a long period, it inhibits the process of
nitrification from ammonium (NH4+) present in high concentration in benthic region [96].
Anoxia also reduces binding of iron to phosphate at the sediment water interface affecting the
availability of phosphorous in water column. Conclusively, blooms decrease the diversity of
benthic forms and influence several trophic levels in food chain. This leads to alterations in
total energy production and decline in the number of secondary producers.

3.4. Loss of Aquaculture

Bloom-induced oxygen depletion, elevated levels of NH4+ and toxins either alone or in
combination cause the death of fish and other aquatic organisms. High pH of waters
persisting during intense cyanobacterial blooms is toxic to certain species of fish [97]. In
temperate regions, hypolimnetic anoxia may destroy the cold water refugia during summer
thus, adversely affecting salmonid fish community.

3.5. Cyanotoxins

Many surface bloom forming cyanobacteria are toxic. Based on morphology, Skulberg et
al. [98] reported that about 40 species of cyanobacteria are toxic, mostly belonging to the
multicellular filamentous genera Anabaena, Nostoc and Oscillatoria. However, the most
prevalent as well as widely studied toxic cyanobacterium is Microcystis aerugenosa.
Cyanobacterial toxins (cyanotoxins) are generally categorized as cytotoxins, hepatotoxins and
86 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

neurotoxins [99] (Table 4). Globally, hepatotoxic freshwater blooms are more common than
neurotoxic blooms [35]. The first report on cyanobacterial toxicity to mammals had come
from Australia [100]. Before this date, the majority of cynotoxicoses cases have been related
to animal poisoning; human causalities are rare [5]. The exact physiological and ecological
basis of toxin production by cyanobacteria is not well known. Cyanobacteria may produce
several variants of a particular toxins as well as group of toxins simultaneously [5].
The toxicity of a water body is not a permanent function, rather it changes with nutrient
status and the collapse of blooms. In general, neurotoxins degrade more rapidly than
hepatotoxins (microcystins) which are more persistent in the system (up to 2 weeks), and may
thrive at high temperatures [59].

Table 4. Different attributes of cyanobacterial toxins

Toxin No. of Chemical structure Biological activity Genus

structural
variants
Microcystins 71 Cyclic heptapeptides Hepatotoxins; inhibitor Anabaena
of protein phosphatases, Anabaenopsis
disturb integrity of Haplosiphon
cytoplasmic membrane, Nostoc
carcinogenes Microcystis
Oscillatoria
Planktothrix
Nodularins 9 Cyclic pentapeptides Hepatotoxins; inhibitor Nodularia
of protein phosphatases,
disturb integrity of
cytoplasmic membrane,
carcinogenes
Cylindrospermopsin 3 Guanidine alkaloid; Necrotic lesions of liver, Anabaena
kidney, spleen, lung, Aphanizomenon
intestine; inhibitor of Oscillatoria
protein synthesis, Phormidium
genotoxic
Anatoxins-a and 5 Alkaloids Inhibitor of acetylcholine Anabaena
Homoanatoxins-a esterase
Anatoxins-a(c ) 1 Alkaloid Inhibitor of acetylcholine Anabaena
esterase Aphanizomenon
Oscillatoria
Phormidium
Saxitoxins 20 Carbamate alkaloids Block sodium channels Anabaena
Aphanizomenon
Cylindrospermopsis
Lyngbya
Planktothrix
Lyngbyatoxins-a 1 Alkaloid Inflammatory agents, Oscillatoria
activates protein kinase Lyngbya
C Schizothrix
Aplisiotoxins 2 Alkaloid Inflammatory agents, Oscillatoria
activates protein kinase Lyngbya
C Schizothrix
Lipopolysaccharides Great Lipopolysaccharides Inflammatory agents, All cyanobacteria
diversity irritate gastrointestinal
tract
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 87

There are a few reports on the relationship between eutrophication and toxin production
[60]. Smith [31] reported that bloom of toxic cyanobacteria is stimulated by P-enrichment.
Contrarily, Boyer et al. [101] reported on a 5-10 fold increase in toxin production in a P-
limited condition rather than that of an N-limited condition. In a eutrophic water bodies, toxic
and non-toxic cyanobacterial species may occur together.

4. CONTROL OF CYANOBACTERIAL BLOOMS

Amongst the various measures used to control cyanobacterial blooms (Figure 2), the most
effective way is nutritional regulation (N and P). Occurrence of cyanobacterial blooms
correlate positively with TN and TP in aquatic systems. Accordingly, the total maximum
daily load (TMDL) guidelines have been developed to forecast cyanobacterial blooms in
aquatic ecosystems. At very high TP (>1mgL-1) and TN (>5mgL-1) small green algae can out-
compete cyanobacteria [30]. Importance of reducing anthropogenic N and P loading in
ecosystem for the control of cyanobacterial bloom formation has long been recognized [102,
103]. N and P are key limiting nutrients in most aquatic and terrestrial ecosystems [104].
However, increased industrialization, agricultural run off containing unused fertilizers,
municipal wastewater and fossil fuel burning has overloaded ecosystems with these nutrients.
N-only reduction has substantially reduced the bloom of non-heterocystous cyanobacteria but
not of heterocystous from. Also, this does not apply in case of brackish and saline waters
[104]. Similarly, in many cases, P-only reduction strategies were unfruitful [104], as P is
rapidly recycled between sediments and waters. Moreover, species like Microcystis can
vertically migrate (buoyancy regulation) to consume phosphorous at the sedimentwater
interface. Studies indicate that no single factor, rather interaction between many factors is
responsible for bloom development [104, 105] (Figure 3). Controlling P input to freshwater
may increase the concentration of N in downstream estuarine and costal waters [104, 106,
107]. Therefore, unless there is clear evidence, control measures focusing exclusively on P or
N reduction would not work [104, 108]. Another major drawback associated with nutrient
control measures is high cost.
The use of chemicals is an effective mean of bloom control. For instance, algicides such
as copper sulphate, sodium penta-chlorophenate, tetra-chlorobenzoquionone, chlorophenyl
dimethyle urea, phenazene1-carboxylic acid, potassium permanganate, 2,3-dichloro-
napthaphenol, dichloropene and chlorine have effectively been used in bloom control. The
exact dose of these algicides depends on pH and other characteristics of water and water
body, and concentration of cyanobacteria. However, efficacy of these chemicals is often short
lived and some of them can have adverse side effects. A suitable algicide should be
inexpensive, potent and specific to the target (i.e., non-toxic to other organisms).
Biological means of bloom control include modification of food web by introducing
suitable organisms that could eat blooms and eaten up by other aquatic organisms such as
fish. Use of cyanophages (LPP-1and 2, SM-1, N-1, AS-1) may help in selective removal of
bloom forming species. The introduction of floating angiosperms such as Lemna, which
shade the water body hence, inhibit the bloom formation. Chrysophytes such as Ochromonas
danica engulf and digest the cells of Microcystis aeruginosa, may also prove fruitful.
88 NK Sharma, KK Choudhary, Rakhi Bajpai et al.

Figure 2. Approaches commonly used for control of blooms.

The Metropolitan Water Board of London has found that artificial circulation of water in
reservoirs is an effective method of reducing bloom development. Manual harvesting (by
nets) of cyanobacterial cells is another method however, small unicels can not be removed by
this method. For them, electrical flocculation method (use of positive current) followed by
routine mechanical removal has been found effective.

30 1000 30 18
Nitrate-N

25
A 25
16 Nitrite-N
Ammonium-N
B
800 14
Chlorophyll a (g.L )

Carbohydrate (g.L )
-1

-1
Protein (mg.L-1)

12
Nitrogen (mg.L )

20 20
-1

600 10

15 15 8

400 6
10 10
4

200 2
5 5
0

0 0 0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month Month

0.8 32 8.4 12

0.7 C PO4
30
D 8.2 11
other phosphorus
Dissolved oxygen (mg.L )
-1

0.6 28 8.0
Temperature (O C)

10
o

26 7.8
0.5 9
P (mg.L-1)

pH

24 7.6
0.4 8
22 7.4
0.3 7
20 7.2
0.2 6
18 7.0

0.1 5
16 6.8

0.0 14 6.6 4
Jan Feb March Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month Month

Figure 3. Seasonal variation in cyanobacterial biomass in relation to environmental and nutritional

factors (Durgakund Pond, Varanasi, India); biomass (A), nitrogen (B), phosphorous (C), temperature
and dissolved oxygen (D); Sharma et al. [104].
 Freshwater Cyanobacterial (Blue-Green Algae) Blooms 89

CONCLUDING REMARKS
Our understanding of freshwater cyanobacterial blooms has improved substantially in
recent times. However, there are many areas still unresolved and require further research.
Some of them are:

1. Impact of nutrient enrichment and climate changes (alone as well as their interaction)
on bloom dynamics (rare abundance species).
2. Impact of hydrological and watershed modifications (e.g., dams and sluices) that
increase residence time on cyanobacterial bloom formation.
3. Analysis of the factors that could be manipulated to control the bloom, even when
environmental conditions are favorable for their development.
4. Interaction/competition between toxic and non-toxic strains and its impact on
cyanobacterial bloom.
5. Impact of physical, chemical and biological factors on the level of toxins.
6. There is a need to understand the effects of cyanobacterial blooms on community
structure and composition rather, than on a particular species.

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