Introduction This is a supplement to the text showing tables. Light Need of light

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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

e-mail 20.satish@gmail.com

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^-2s^-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

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