1
Factors controlling induction of reproduction in algae – review: A Supplement to the text
S. C. Agrawal
Department of Botany, University of Allahabad, Allahabad 211002, India
Introduction
This is a supplement to the text showing tables.
Light
Need of light
A period of illumination with visible light of certain intensities above a particular minimal threshold value and below a particular high level was required for the initiation and maturation of reproductive cells in members of all major groups of algae (Table 1). In most of the cases in which algae are known to release zoospores or gametes in response to changes in light, release occurred at daybreak (or artificial day in culture on a light/ dark cycle, Table 1).
Table 1 Visible light was stimulatory to algal reproduction
Alga Structure / process Light/ favorable light Reference
induced intensity
Cyanophyceae
Anabaena doliolum Akinete White light (10- Pandey and Kashyap
60 μmol photons m-2 s-1) 1987
Anabaena iyengarii Akinete White light (> 6 Agrawal and Singh
μmol photons m-2 s-1) 2000
Aphanothece nidulans Cell division White light (40 Agrawal and Pal
μmol photons m-2 s -1) 2003
Apanothece pallida Cell division White light (40 Gupta and Agrawal
μmol photons m-2 s-1) 2008
Chroococcus minor Cell division White light (40 Agrawal and Pal
μmol photons m-2 s-1) 2003
Cylindrospermopsis Akinete White light (100 Moore et al. 2005
raciborskii μmol photons m-2 s-1)
Gloeocapsa Cell division White light (40 Agrawal and Pal
aeruginosa μmol photons m-2 s-1) 2003
Gloeocapsa Cell division White light (40 Gupta and Agrawal
atrata μmol photons m-2 s-1) 2008
Nostochopsis Akinete White light Agrawal and Singh
lobatus (> 6 μmol photons m-2 s-1) 2000
Westiellopsis Akinete White light Agrawal and Singh
prolifica (> 2 μmol photons m-2 s-1) 2000
Chlorophyceae
Basicladia crassa Release of Visible light Hamilton 1948
flagellated cells
Caulerpa sp. Spore Visible light Suto 1950
Chara fragilis Sex organs Visible light Karling 1924
Chlamydomonas Gametogenesis, White light Moewus 1933,
eugametos mating (during N- 1934; Lewin
starvation) 1954; Förster and
Wiese 1954;
Förster 1957;
Stifter 1959;
Lorch and
Karlander 1973
Chlamydomonas Mating of Visible light Lewin 1956;
moewusii gametes (in absence of Förster 1959
CO2)
Chlamydomonas Mating of Visible light Van Winkle- Swift
monoica gametes and Bauer 1982
Chlorella Autospore White light Agrawal and
vulgaris mother cell (> 10 μmol Singh 2001
photons m-2 s-1)
Closterium Cell division, Visible light Hogetsu and
ehrenbergii Cell aggregation (and nitrogen Yokoyama
during conjugation depletion) 1979
Closterium Cell division Visible light Kato et al.
peracerosum linked to (> 60 μmol photons m-2 s-1) 1983
strigosum conjugation
littorale
Cosmarium Initial phase Visible light Starr 1954 c;
botrytis of conjugation 1955
Enteromorpha sp. Zoospore induction Visible light Lersten and Voth
1960
Enteromorpha Swarmer Visible light Dan et al. 2002;
prolifera release/ asexual (> 16 μmol photons Fu et al. 2008
reproduction m-2 s-1, 40 μmol
photons m-2 s-1)
Eudorina elegans Sperm packets Increased light Szostak et al.
intensity 1973
Halicystis ovalis Gamete liberation Shortly after Hollenberg
Sun rise 1936
Hormidium flaccidum, Zoospore Visible light Strasburger 1878;
H. nitens liberation Gerneck 1907;
Vischer 1933
Hydrodictyon Zoospore formation Visible light Neeb 1952
reticulatum
Daughter net White light Agrawal and
(40 μmol photons m-2 s-1) Pal 2003
Micrasterias Zygote White light Imaizumi and
thomasiana (30- 60 μmol photons m-2 s-1) Doida 1990
Netrium digitus Sexuality Visible light Biebel 1964
Oedogonium Gametogenesis Visible light Starr 1960;
cardiacum, (16: 8 h LD cycle) Hoffman 1960;
Oedogonium sp. Hill and Machlis
1970; Coss and
Pickett- Heaps
1973; Agrawal
and Pal 2003
Pandorina unicocca Mating Visible light Rayburn 1974
Pithophora Akinete White light Chaudhary and
oedogonia (40- 70 μmol photons m-2 s-1) Singh 1987;
Agrawal and
Sarma 1983;
Agrawal and
Singh 2000
Protosiphon Cyst formation, Bright sun light Strasburger 1878;
botryoides Zoospore liberation Klebs 1896;
Vischer 1933;
Gerneck 1907
Rhizoclonium Zoospore formation, White light Gupta and
hieroglyphicum liberation (40 μmol photons Agrawal 2004 b
m-2 s-1)
Scenedesmus Gametogenesis Visible light Cain and Trainor
obliquus (but can occur 1976
in absence of
light)
Schizomeris sp. Zoospore formation Visible light Kulfinski and
(darkness is Henschel 1969
inhibitory)
Spirogyra spp. Conjugation Visible light Pessoney 1968;
Simons et al.
1984
Stigeoclonium Akinete White light Agrawal and
pascheri (60- 70 μmol photons m-2 s-1) Sarma 1982 a
Ulothrix zonata Zoosporogenesis Minimal at 32 Graham et al 1985
μmol photons m-2 s-1
Ulva fasciata Swarmer discharge Visible light Mohsen et al.
(2- 40 μmol 1973 b
photons m-2 s-1)
Ulva lactuca Sporogenesis, White light Kalita and
(=U. fenestrata) Gametogenesis (40 μmol Tytlianov 2003
photons m-2 s-1)
Ulva mutabilis Sporulation Visible light Nordby 1977
(lower limit 3- 5
μmol photons m-2 s-1,
optimum level
60 μmol photons m-2 s-1)
Ulva pertusa Sporulation Blue or red light Hiraoka and
(60- 200 μmol photons Enomoto 1998;
m-2 s-1, 12: 12 h Han et al. 2003;
LD cycle) Han and Choi 2005
Ulva Zoospore Visible light Hiraoka et al. 2003
spinulosa (50 μmol photons m-2
s-1, 12: 12 h
LD cycle)
Xanthophyceae
Botrydium Zoospore Visible light Strasburger
granulatum liberation 1878
Vaucheria Zoospore Visible light Strasburger
repens liberation 1878
Vaucheria Sex organs Visible light League and
sessilis Greulach
1955;
Starr 1960
Bacillariophyceae
Chaetoceros sp. Spore formation Visible light Hargraves and
French 1975;
Hollibaugh et al.
1981
Cocconeis scutellum Sexual Visible light Mizuno and
reproduction (60 μmol photons m-2s-1) Okuda 1985
Coscinodiscus Sexuality, Visible light Werner 1971;
asteromphalus, auxospore (prolongation Roshchin and
C. granii of photoperiod Lutsenko 1972
and light
intensity)
Eunotia Resting spore Visible light von Stosch and
soleirolii (ca. 30 μmol photons Fecher 1979
m-2 s-1)
Lithodesmium Sexuality, Stronger Manton et al.1968
undulatum auxospore visible light
Melosira Sexuality, Visible light Bruckmayer-
nummuloides auxospore Berkenbusch
1954
Nitzschia Gametes, Increased with Davidovich
lanceolata auxospore total light energy 1998
Pseudo- nitzschia Gametes Visible light Hiltz et al. 2000
multiseries (100 μmol photons
m-2 s-1, 16: 8
h LD cycle)
Rhabdonema Sexuality Visible light Rozumek 1968
adriaticum
Stephanopyxis Gametes, Visible light Steele 1965;
palmeriana auxospore (8- 70 μmol photons m-2 s-1) Drebes 1966
Stephanopyxis Sexuality Visible light von Stosch and
turris Drebes 1964
Thalassiosira Spermatogenesis Visible light Vaulot and
weiwssflogii Chisholm 1987
Rhodophyceae
Acrochaetium Tetrasporangia Visible light West 1971
proskaueri (30 μmol photons m-2 s-1)
Bangia Monospore Visible light Graham and
atropurpurea Graham 1987
Champia Fertilization Visible light Thursby et al.
parvula 1985
Polysiphonia Gametes, Visible light Edwards 1970
denudata tetraspores
Scinaia confusa Tetrasporangia Visible light Ramus 1969
(=Pseudogloio- (15- 20˚C,
phloea confusa) N- depletion)
Phaeophyceae
Desmarestia Oogonia, Visible light Chapman and
aculeata antheridia Burrows 1970
Fucus Gametes In light in Serrão et al.
vesiculosus release natural population 1996
Halosiphon Oogonia, Strong white Maier 1984
tomentosus antheridia fluorescent light
(=Chorda tomentosa)
Laminaria Oogonia, Visible light Kain 1964
hyperborea antheridia
Macrocystis Egg, sperm Visible light Lüning and
pyrifera (> 5 μmol photons m-2s-1) Neushul 1978;
Deysher and
Dean 1984
Nereocystis Antheridia, Visible light Vadas 1972
luetkeana oogonia
Sargassum sp. Spore Visible light Suto 1950
Zonaria farlowii Spore, propagule Visible light Liddle 1968
Dinophyceae
Peridinium bipes Cyst Increased Park and
illumination Hayashi 1992,
(at 15- 20˚C) 1993
Pyrophacus Cyst Visible light Zonneveld and
steinii (70 μmol photons m-2 s-1) Susek 2007
at 27˚C
Darkness or light limitation
A reduction in the availability of light or transfer from light to darkness under laboratory conditions have been shown to stimulate akinete formation in some blue- green algae, zoospore formation in many green and yellow- green algae, egg release in some brown algae and cyst or resting- cell formation in diatoms Stephanopyxis turris and Coscinodiscus wailesii and in a dinoflagellate Scrippsiella hangoei (Table 2). Illumination with white light inhibited zoospore formation strongly possibly by inhibiting cytoplasmic cleavage in parent cells. Cessation of growth due to light or energy limitation marked the beginning of akinete differentiation in Anabaena circinalis, A. cylindrica, Nostoc punctiforme and Nostoc strain PCC 7524 (Table 2). The cultures in which N. punctiforme had differentiated into akinetes contained approximately 10- fold higher concentrations of 2-methyl hopanoids than did cultures that contained only vegetative cells (Meeks et al. 2002; Doughty et al. 2009).
Table 2 Darkness or light limitation induced reproduction in some algae
Alga Structure/ Triggering factor Reference
process induced
Cyanophyceae
Anabaena Akinete Light limitation in Fay et al. 1984
circinalis a dense filament
suspension
Anabaena Akinete A reduction in the Fogg 1949;
cylindrica availability of Wolk 1965;
light Nichols et al.
1980
Nostoc punctiforme Akinete Insufficient light Meeks et al.
(or P limitation) 2002;
Doughty et al.
2009
Nostoc strain Akinete Light or energy Sutherland
PCC 7524 limitation et al. 1979
Chlorophyceae
Botryococcus Zoospore, Darkness Pavel and
sudeticus gamete Vladislav 2007
emergence
Chlamydomonas Zoospore Middle of the Mergenhagen
reinhardtii formation dark period 1980
Chlorococcum Gamete Darkness O’Kelley 1983
echinozygotum formation,
liberation
Cladophora sp. Zoospore Low light, short Mason 1965;
formation days (13- 20˚C, Pantastico and
vitamin limitation) Zenaida 1973;
Hoffman and
Graham 1984
Gloeocystis sp. Zoospore Darkness Gerneck 1907
formation
Hormidium Zoospore Darkness Klebs 1896;
flaccidum, formation Gerneck 1907
H. nitens,
Hormidium sp.
Planophila sp. Zoospore Darkness Gerneck 1907
formation
Protosiphon Zoospore Dim light, yellow Klebs 1896;
botryoides formation light, or darkness Williamson
1962; Stewart
and O’Kelley
1966; Durant
et al. 1968
Gamete Darkness or Maher 1946,
formation diffuse light 1947
Stigeoclonium Zoospore Darkness or Agrawal
pascheri formation 5 μmol photons m-2 s-1 1993
of white light
Ulva flexuosa Zoospore Darkness of Kolwalkar
(=Enteromorpha formation 1- 7 days et al. 2007
flexuosa)
Ulva Gamete An obligatory Lüning et al.
pseudocurvata release dark period of 2008
at least 1 h
before gamete
release early in
the morning
Xanthophyceae
Botrydium Zoospore Darkness Klebs 1896
granulatum formation
Ophiocytium sp. Zoospore Darkness Gerneck 1907
formation
Vaucheria Zoospore Darkness Klebs 1896
repens formation
Bacillariophyceae
Coscinodiscus Resting cell Darkness Nagai and Imai
wailesii formation 1999
Stephanopyxis Cyst Decreasing light von Stosch
turris formation intensity and and Drebes
photoperiod 1964
(decreasing
temperature)
Phaeophyceae
Halosiphon Egg release Dark period Maier 1984
tomentosus
(=Chorda tomentosa)
Saccharina japonica Egg release Dark period Tseng et al.
(=Laminaria japonica) 1959
Saccharina Egg release Dark period Lüning 1981 b
latissima
(=Laminaria saccharina)
Silvetia compressa Gamete liberation Darkness Jaffe 1954
(=Pelvetia fastigiata)
Dinophyceae
Scrippsiella hangoei Cyst formation Dark conditions Rintala et al.
2007
Light quality
Red light favored akinete formation in most of the blue- green algae assessed, and both red and blue light the reproduction in different green algae assessed (Table 3). Blue light dominate in sublittoral clearest oceanic water (Jerlov 1976) and it also support reproduction in several brown algae, in diatoms Biddulphia and Ptychodiscus, and in a red alga Nitophyllum investigated (Table 3). Blue and green light penetrates deep water, while red light is available in surface water.
Table 3 Light quality favorable for induction of algal reproduction
Alga Structure/ Favorable light Reference
process induced quality
Cyanophyceae
Anabaena circinalis Akinete Red light Thompson et al.
2009
Anabaena doliolum Akinete Yellow light Pandey and
Kashyap 1989
Anabaena sp. Akinete Red light Kaushik and
Kumar 1970
Anabaena torulosa Akinete Red light Fernandes and
(Incandescent Thomas 1982
light)
Fischerella sp. Akinete Red light Kaushik and
Kumar 1970
Gloeotrichia Akinete Green light Wyman and
echinulata ( in absence Fay 1986
of combined
nitrogen)
Nodularia spumigena Akinete Red light Pandey and
Talpasayi 1980,
1982
Westiellopsis Akinete Blue and Agrawal and
prolifica Green light Sharma 1994
Chlorophyceae
Acetabularia Cap Small quantity Beth 1953;
crenulata of blue light in Terborgh 1965;
red light Terborgh and
Thimann 1965
Bryopsis plumosa Gamete release Blue light Mine et al. 1996
Chlamydomonas Gametogenesis, Blue light Sager and Granick
eugametos, mating 1954; Förster 1957,
C. reinhardtii, 1959; Treier et al.
C. moewusii 1989; Weissig and
Beck 1991; Beck
and Haring 1996;
Hegemann et al.
2001
Chloromonas sp. - D Reproduction Blue light Hoham et al. 2000
Cladophora sp. Zoospore Both blue and Cook and Price
liberation and red light 1928
Akinete Red light Pantastico and
formation Zenaida 1973
Golenkinia sp. Spermatogenesis, Both blue and Ellis and Machlis
sexuality red light 1968
Monostroma sp. Gamete Blue light Shihira 1958
liberation
Pithophora oedogonia Akinete Blue light Agrawal and
Sarma 1983
Protosiphon Zoospore Yellow light Durant et al. 1968
botryoides production
Stigeoclonium Zoospore, Red light Abbas and
amoenum zygote formation Godward 1963
Stigeoclonium Akinete Red light Agrawal and
pascheri Sarma 1982 a
Trebouxia sp. Aplanospore Red light Giles 1970
Ulva pertusa Sporulation Blue or red light Han et al. 2003
Bacillariophyceae
Biddulphia sp. Auxospore Blue and green light Baatz 1941
Haslea ostrearia Auxospore Red light Mouget et al. 2009
Ptychodiscus brevis Gametes, Zygotes Blue light Walker 1982
Rhodophyceae
Nitophyllum punctatum Tetraspore liberation Blue light Sagromsky 1961
Phaeophyceae
Dictyota dichotoma Reproduction Red light Müller and Clauss
1976
Laminaria farlowii Gametophytes Blue light Lüning and
Neushul 1978
Macrocystis Eggs and sperms Blue light Lüning and
pyrifera production Neushul 1978;
Deysher and
Dean 1984
Petalonia fascia, Zoospore formation Blue light Hsiao 1970;
P. zosterifolia and liberation Lüning and
Dring 1973
Pterygophora Gametophytes Blue light Lüning and
californica Neushul 1978
Saccharina japonica Zoosporangial sori, Blue light Cuijuan et al.
(=Laminaria zoospore release, 2005; Shi et al.
japonica) gametophytes 2005; Mizuta
et al. 2007
Saccharina latissima Eggs and sperms Blue light Hsiao and Druehl
(=Laminaria production 1971; Lüning and
saccharina) Dring 1972, 1975;
Lüning 1980 a
______
Photoperiod
Short days induced sexual reproduction in many red and brown algae, sexuality in diatoms Cocconeis scutellum, Coscinodiscus concinnus and Navicula ostrearia and in dinoflagellate Gymnodinium pseudopalustre, cyst formation in some dinoflagellates, formation of zoospores in Cladophora glomerata, aplanospores in Dunaliella salina and sporulation in Monostroma sp. (Table 4).
Table 4 Photoperiod (day length) responsible for algal reproduction.
Alga Structure/ Photoperiod Reference
process induced (day length)
responsible
for induction
Chlorophyceae
Chloromonas sp. - D Reproduction Long days Hoham et al.
(20: 4 h LD 2000
cycle)
Cladophora glomerata Zoosporogenesis Short days Hoffman and
(8: 16 h LD) Graham 1984
Derbesia Sporangia Long days Hustede 1964
neglecta
Derbesia tenuissima Sporangia Both long and Ziegler and
Short days Kingsbury 1964;
Rietema 1973
Dunaliella salina Aplanospore Short days Borowitzka and
(cool temperature) Huisman 1993
Klebsormidium Zoospore Diurnal regime of Cain et al.1974
flaccidum 18 h light and 16 h
dark
Micrasterias Zygote Long days Imaizumi and
thomasiana (> 12 h light Doida 1990
per day)
Monostroma sp. Sporulation Short days Lüning 1980 b,
(8: 16 h LD), 1981 a
(5- 15˚C)
Ulothrix zonata Zoosporogenesis Either short days Lôkhorst and
(8: 16 h LD) or Vroman 1972;
long days (16: 8 h 1974 a, b
LD)
Gametogenesis Long days (16: 8 h Graham et al.
LD) 1985
Ulva mutabilis Meiotic zoospore Long days Nordby 1977
(15 h light per day)
Bacillariophyceae
Cocconeis scutellum Sexuality Short days Mizuno and
(10: 14 h LD) Okuda 1985
Coscinodiscus Auxospore Short days Holmes 1966
concinnus
Haslea ostrearia Sexuality Short days Mouget et al.
(6- 10 h light 2009
per day)
Navicula ostrearia Auxospore Short days Neuville and
(6 h light per day) Daste 1975
Stephanopyxis Gametes Long days Steele 1965;
palmeriana (15- 21˚C) Drebes 1966
Rhodophyceae
Acrochaetium Tetrasporangia Short days Abdel- Rahman
asparagopsis 1982
Acrochaetium Tetrasporangia Short days West 1968
pectinatum (6- 10 h light
per day)
Acrochaetium Monospore, Short days West 1971
proskaueri tetraspore (8: 16 h LD)
Acrosymphyton Tetrasporangia Short days Cortel- Breeman
purpuriferum (8: 16 h LD) and Hoopen 1978
Asparagopsis armata Tetrasporangia Short days Oza 1976, 1989;
(Falkenbergia- phase) (15- 17˚C, Guiry and Dawes
reduced N and 1992
P, presence of
iodine)
Bangia fuscopurpurea Conchospore Short days Richardson and
(< 12 h light Dixon 1968;
per day) Richardson 1970
Monospore Long days Conway and
(16 h light per Cole 1977
day, summer
temperature)
Bonnemaisonia Tetrasporangia Short days Chihara 1961;
hamifera (< 12 h day Chen et al.
length) 1970 a; Lüning
1980 b, 1981 a
Calosiphonia Monosporangia Long days Mayhoub 1973,
vermicularis 1975
Tetrasporangia Short days Mayhoub et al.
1976
Chondrus giganteus Gametangia, Short days Brodie et al.
tetrasporangia 1994
Chondrus nipponicus Gametangia Short days Brodie et al. 1991
Chondrus ocellatus Gametangia Short days Brodie et al. 1993
Chondrus pinnulatus Tetrasporangia Short days Brodie and Guiry
1988
Constantinea subulifera Tetrasporangia Short days Powell 1964
Dumontia contorta Sexuality Short days Rietema and
Breeman 1982
Eupogodon planus Gametangia Both short and Orfanidis et al.
long days 1999
Tetrasporangia Short days Orfanidis et al.
1999
Gigartina acicularis Gametangia Short days Guiry and
(14- 18˚C) Cunningham
1984
Gigartina franciscana Tetraspore Short days West 1972 b
(Petrocelis) (8: 16 h LD)
Hyalosiphonia caespitosa Tetrasporangia Short days Umezaki 1972
(7: 17 h LD)
Kylinia rosulata Monosporangia Short days Stegenga and
Tetrasporangia Long days Van Wissen
1979
Meredithia microphylla Tetrasporangia Short days Guiry and Maggs
(15˚C) 1984
Porphyra carolinensis Conchospore Short days Wilson and
release (low temperature) Kapraun 1986
Porphyra miniata, Monospore Long days Chen et al. 1970b;
P. elongata (summer Conway and Cole
(=P. rosengurttii) temperature) 1977; Kapraun
and Luster 1980
Porphyra tenera Monospore Long days Iwasaki 1961
(summer
temperature)
Conchosporangia Short days Kurogi and Sato
(8: 16 h LD), 1962;
(15˚C) Dring 1970
Scinaia confusa Tetrasporangia Long days Ramus 1969
(=Pseudogloiophloea (16: 8 h LD)
confusa)
Rhodochorton Tetrasporangia Short days West 1972 a;
purpureum (10 ˚C) Dring and West
1983
Thuretellopsis Tetrasporangia Short days Richardson and
peggiana (9: 15 h LD) Dixon 1970
Phaeophyceae
Desmarestia tabacoides Sex organs Short days Nakahara and
Nakamura 1971
Fucus distichus Receptacles Short days Bird and
(< 12 h light) McLachlan 1976
Saccharina longicruris Gametogenesis Short days Byrne and
(=Laminaria longicruris) Craigie 1971
Scytosiphon lomentaria Zoospore Short days Lüning 1981 a
release
Sphacelaria rigidula Gametangia Long days Hoopen et al. 1983
Undaria pinnatifida Fertility of Short daylength Choi et al. 2005
gametophyte (8 h)
Dinophyceae
Gymnodinium Gametes Short days von Stosch 1973
pseudopalustre (10 h light),
(15˚C)
Scrippsiella Cyst Short days Sgrosso et al 2001
trochoidea, (of different photoperiod
S. rotundata, regimes)
S. operosa,
Pentapharsodinium
tyrrhenicum
Temperature
Blue- green algae formed akinetes in cultures at water temperatures of 10 to 37˚C depending upon the species (Table 5). Most of the green algae investigated reproduced in cultures optimally at 20- 25˚C, but Scenedesmus obliquus and Ulva fenestrata formed gametes at 15˚C and Urospora sp. at 4˚C (Table 5). Different diatoms showed sexuality and resting spore formation at 2- 30˚C. Temperatures most favorable for reproduction of red algae assessed were 10- 20˚C, but subarctic Clathromorphum formed conceptacles at 2- 3˚C. Reproduction in brown algae occurred at somewhat lower temperatures than in red algae assessed (Table 5). In a brown alga, different life cycle stages may occur in different seasons since they have different temperature optima. In kelp members (Halosiphon=Chorda, Macrocystis), gametogenesis occurred at somewhat lower temperatures than zoosporogenesis. Dinoflagellate Alexandrium tamarense (= Gonyaulax tamarensis) formed sexual stages at 10- 12˚C and Gymnodinium pseudopalustre formed gametes at 15˚C (Table 5). Chrysophyceae members formed statospores at temperatures less than 20˚C and a Raphidophyceae member Gonyostomum formed resting cysts at temperatures less than 10˚C (Table 5).
Table 5 Optimum temperature (˚C) to induce reproduction in different algae
Alga Structure/ Optimum Reference
process induced temperature
(˚C)
Cyanophyceae
Anabaena crassa, Akinete 10 and 15 Li et al. 1997
A. flos-aquae,
A. mucosa, A. spiroides,
A. ucrainica
Anabaena doliolum Akinete 32 ± 2 Pandey and
(also growth) Kashyap 1987
Anabaena torulosa Akinete 37 > 32 > 29 Fernandes and
Thomas 1982
Cylindrospermopsis Akinete 15- 25 Moore et al. 2003,
raciborskii 2005
Nodularia spumigena Akinete 32 Pandey and