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Table 7 Inorganic substances and nutrients stimulatory to algal reproduction in culture

Alga or flora Structure/ Stimulatory Reference

process induced chemical

Cyanophyceae
Anabaena iyengarii Akinete Nitrogen and Agrawal and

phosphorus Misra 2002

compounds
Anabaena spp. Akinete Nitrate,sulfate, Canabaeus 1929;

sodium chloride Kanta and Sarma

1980
Anabaena Akinete Ammonium Sarma and Garg

torulosa carbonate, 1985, Sarma et al.

bicarbonate, 2004

nitrate
Calothrix parietina Hormogonium Excess phosphate Wood et al.1986
Calothrix sp. Hormogonium Iron Douglas et al. 1986
Cylindrospermopsis Akinete Phosphorus Moore et al. 2003,

raciborskii (22- 75 μg/L) 2005
Gloeotrichia Akinete Iron, magnesium, Sinclair and

echinulata calcium, sulfate Whitton 1977
Nodularia Akinete Ammonium Pandey and

spumigena carbonate, nitrogen Talpasayi 1980;

and carbon Pandey 1989;

compounds Myers et al. 2011

(higher irradiance,

low and high salinities)

Nostoc PCC Akinete Presence of Sutherland et al. 1979;

7524 nitrogen source Ahluwalia and

Kumar 1983
Nostochopsis Akinete Nitrate, Agrawal and

lobatus phosphate, Misra 2002

other nutrients

of BG 11 medium
Scytonema Akinete Ammonium nitrate Sarma and Ghai

fritschii 1998
Westiellopsis Akinete Sodium chloride, Agrawal and

prolifica magnesium sulfate, Sharma 1994;

nitrate, phosphate, Agrawal and

other nutrients of Misra 2002

Bold’s basal medium



Chlorophyceae
Chaetophora sp. Zoosporogenesis Phosphorus, Gibson and

nitrogen Whitton 1987


Chlamydomonas Mating, Calcium, Wiese and Jones

eugametos flagella magnesium 1963

agglutination


Chlamydomonas Mating Nitrate, calcium Lewin 1954;

moewusii Bernstein and

Jahn 1955;

Trainor 1959

Chlorella Autospore Calcium Carroll and Stiller

pyrenoidosa release 1975

(at low pH)


Chlorococcum Zoospore Calcium Gilbert and

echinozygotum release O’Kelley 1964

Chlorococcum Sexuality NaCl, MgCl2 Hardy et al. 2000

hypnosporum
Cladophora Zoosporangium Nitrogen, Agrawal and

glomerata formation, Phosphorus, Misra 2002

zoosporogenesis other nutrients


Closterium Conjugation Carbon dioxide Lippert 1973

ehrenbergii,

C. moniliferum
Coelastrum Daughter Calcium salts Rayss 1915

proboscideum coenobia (0.25- 1.75 %)
Cosmarium Pairing of Carbon dioxide Starr 1955, 1964

botrytis conjugants,

conjugation


Draparnaldia Zoosporogenesis Phosphorus, Gibson and

sp. nitrogen Whitton 1987


Golenkinia Mating, Calcium Ellis and Machlis

minutissima zygospore (1 mmol/L) 1968
Hydrodictyon sp. Gamete formation Phosphate Neeb 1952
Mesotaenium Conjugation Calcium Tiftickjian and

kramstai (> 0.5 mmol/L), Rayburn 1986

magnesium

(> 1.0 mmol/L)


Micrasterias Zygote Carbon dioxide Imaizumi and

thomasiana in nitrogen- free Doida 1990

Waris medium


Oedogonium Oogonia, Soil water medium Starr 1960;

cardiacum, antheridia containing CaCO3 Hill and Machlis

O. rupestre and 5 % solution 1970; Coss and

of NaHCO3 Pickett- Heaps 1973


Oedogonium Zoosporogenesis Carbon dioxide Gussewa 1931,

cardiacum, Retallack and

O. capillare von Maltzahn

1968
Pandorina Mating, Calcium chloride, Rayburn 1974



unicocca zygote nitrate

strains 105- 106


Protosiphon Coenocyst Nitrate Maher 1947

botryoides

Zoospore Sulfate and Nitrate O’Kelley and

formation, (for zoospore Herndon 1959,

release formation), 1961; O’Kelley

Calcium (for and Deason

zoospore release) 1962


Rhizoclonium Zoosporangium Nitrogen, Agrawal and

hieroglyphicum and zoospore phosphorus, Misra 2002;

formation, other nutrients Gupta and

zoospore release of Bold’ s basal Agrawal 2004 b

medium
Staurastrum sp. Conjugation Carbon dioxide Winter and Biebel

1967
Stigeoclonium Akinete Magnesium sulfate, Agrawal and

pascheri nitrate, phosphate Sarma 1982 b
Stigeoclonium sp. Zoospore Phosphorus, Gibson and Whitton

nitrogen 1987


Stigeoclonium sp. Reproduction Carbon dioxide Abbas and

Godward 1963


Volvox sp. Gonidia Carbon dioxide Starr 1970 b
Volvulina steinii Mating Calcium Carefoot 1966
Zygnema Conjugation Nitrate in proper Jost 1953

circumcarinatum levels


Xanthophyceae
Botrydiopsis sp., Zoosporogenesis, Nitrogen, calcium Pecora and

Bumilleriopsis sp., zoospore release Rhodes 1973

Pseudollumilleri-

opsis sp.

Bacillariophyceae
Coscinodiscus Sexuality Salinity increase Jensen et al.

wailesii (5 %) 2003
Cyclotella cryptica, Sexuality Increase in sodium Schultz and

C. meneghiniana concentration Trainor 1968
Licmophora Sexuality Extra addition of Kumar 1978

hyalina sodium nitrate

(7.5 mg/ 10 mL)



Phaeophyceae
Alaria Zoosporangial Higher nitrogen Kumura et al. 2006

crassifolia sori and phosphorus

levels
Dictyota Reproduction Boron Nasr and Bekheet



dichotoma 1970


Ectocarpus Reproduction Iodine Woolery and

siliculosus Lewin 1973


Fucus evanescens Embyo Boron McLachlan 1977

(=F. edentatus) development



Macrocystis Oogonia Increased nutrient Muñoz et al.

pyrifera concentration 2004
Petalonia Reproduction Iodine Hsiao 1969

fascia
Saccharina Oogonia Nitrate, phosphate Hsiao and Druehl

latissima (limited quantities) 1973 a

(=L. saccharina)


Undaria Zoosporangial Nitrogen, Kumura et al. 2006 pinnatifida sori phosphorus

Dinophyceae
Many Gametes Nutrients- replete Garcés et al. 2002

Dinoflagellates waters



Chrysophyceae
Dinobryon Encystment Phosphate Sandgren 1981

cylindricum

Organic substances inducing reproduction

Organic carbon compounds like acetate, citrate, fumarate; monosaccharides, oligosaccharides or polysaccharides (glucose, fructose, glucuronate or sugar); organic acids and alcohol at limited levels can act as a source of carbon and energy to induce reproduction in some algae (Table 8). Amino acids (tryptophan, tyrosine, aspartic acid, arginine, glutamic acid) at some low levels can be a source of carbon and nitrogen to some algae and induced reproduction in them (Table 8). Amino acid analogues like canavanine (an arginine analogue) and mixture of amino acids analogues (methionine- dl sulfoximine, hydroxylsine, thienyl- dl- alanine, azatryptophan) favored reproduction in some algae at low levels (Table 8) but amino acid serine at 100- 600 ppm inhibited akinete formation in Stigeoclonium pascheri (Agrawal 1988a). Thus some amino acids at suitable low levels were helpful to reproduction of some algae. Drugs (aminophylline and theophylline), cAMP and dibutyryl cAMP promote flagella agglutinability and mating in Chlamydomonas (Table 8).

Many algae required vitamins for survival and growth (auxotrophy). A limited levels of vitamin B1 (thiamine) and B12 (cyanocobalamine) induced zoosporogenesis in Cladophora glomerata, and of vitamine C (ascorbic acid) the auxospore formation in Melosira nummuloides. In Dinobryon cylindricum, encystment was induced by vitamin B1, but sexual reproduction was inhibited by vitamin B1, B7 (biotin) and B12 (Sandgren 1981). Vitamin B1, B2 and C at high levels inhibited akinete formation in Pithophora oedogonia (Agrawal 1988b) and vitamin C at high levels the akinete formation in Stigeoclonium pascheri (Agrawal 1988a). Only low levels of some vitamins may be helpful to reproduction of some algae.

Plant growth hormones IAA, GA3 and kinetin at suitable low concentrations induced reproduction in some algae (Table 8). IAA and GA3 at 0.01- 500 ppm decreased akinete formation in Pithophora oedogonia (Agrawal 1984). Only limited levels of some plant hormones can be useful to reproduction in algae. Exact role of plant hormones on algal reproduction is not known.



Table 8 Organic substances inducing reproduction in algae

Alga or flora Structure/ Organic substance Reference

process induced responsible

________________________________________________________________________



Cyanophyceae
Anabaena Akinete Organic acids, Fay et al. 1984

circinalis ethanol (1 %),

fumarate, citrate,

acetate
Anabaena Akinete Canavanine (an Wolk 1965;

cylindrica arginine analog), Nichols et al.

Neo- peptone, 1980; Sharma

some polypeptides, 1984; Hori

sodium acetate, et al. 2003

calcium glucuronate
Anabaena Akinete Glucose Tyagi 1974

doliolum
Anabaena Akinete Tryptophan, tyrosine Ahluwalia 1985

oscillarioides aspartic acid
Anabaena spp. Akinete Acetate, citrate Canabaeus 1929;

Kanta and Sarma

1980
Anabaena Akinete Acetate, citrate, Sarma and Garg

torulosa fructose 1985; Sarma

et al. 2004


Cylindrospermum Akinete Tryptophan, Hirosawa and

licheniforme Glucuronate Wolk 1979 a
Nodularia Akinete L- aspartic acid, Pandey and

spumigena L- arginine Talpasayi 1980
Nostoc Akinete Fructose in Summers et al. 1995

punctiforme darkness

(a mutant strain)




Chlorophyceae
Acetabularia sp. Caps IAA, kinetin Spencer 1968;

(at lower levels) Driessche 1984


Chlamydomonas Mating, Dibutyryl- cAMP, Pijst 1985;

eugametos zygote c-AMP, Tomson et al. 1985;

formation aminophylline, Kooijman et al.

theophylline 1990
Chlamydomonas Agglutinability Dibutyryl- cAMP Greenwood 1973;

reinhardtii of gametes Goodenough 1989

Chlamydomonas Sexuality Kinetin, abscisic Ishiura 1976

sp. in dark acid (at lower levels)


Cladophora Zoosporangium Vitamin B1, B12 Hoffman and

glomerata formation, Graham 1984

zoosporogenesis




Derbesia Spore IAA at lower Hustede 1964

neglecta levels
Micrasterias Zygote Amino acid analogs Imaizumi and

thomasiana 1- methionine- dl Doida 1990

sulfoximine (MSO),

Ō- hydroxylsine (Hyl),

B- 2- thienyl- dl- alanine,

dl- 7- azatryptophan
Oedogonium hatei Oogonia IAA, GA3 at lower Singh and

levels Chaudhary 1988


Protosiphon Zygote Sucrose (1 %) Maher 1947

botryoides


Rhizoclonium Zoosporangia Urea (200 ppm) Gupta and

hieroglyphicum dehiscence Agrawal 2004 b
Spirogyra sp. Parthenospore Sugar Klebs 1896
Stigeoclonium Akinete IAA Agrawal and

pascheri (0.1- 1 ppm), Sarma 1984

GA3

(10- 500 ppm)
Trebouxia sp. Aplanospore IAA at lower levels Giles 1970
Trentepohlia Reproduction Auxins Jose and

effusa Chowdhary 1978
Ulothrix sp. Zoospore IAA, GA3 at Conrad et al. 1959

lower levels


Ulva mutabilis Sporulation Nicotinic acid Thiadens and

(45 μg/ 50 mL), Zeuthen 1967



d-panthothenate,Ca

(37.5 μg/ 50 mL),

Thiamine- HCl

(25 μg/ 50 mL),

Biotine

(0.05 μg/ 50 mL),



Riboflavine

(22.5 μg/ 50 mL),

Dithioctic acid

( 0.5 μg/ 50 ml),

Pyridoxamine 2HCl

(2.5 μg/ 50 mL),

Pyridoxal- HCl

2.5 μg/ 50 mL)


Volvox sp. Sexual gonidia Glutamic acid, Starr 1970 b;

L- tryptophan Starr et al. 1980;

Miller and Starr

1981; Jaenicke

1982

Xanthophyceae
Vaucheria sessilis Zoospore IAA at suitable von Denffer and

concentration Hustede 1955;

Hustede 1957

Bacillariophyceae
Coscinodiscus Spermatogenesis Methanol (40 mg/L) Werner 1971

asteromphalus
Melosira Auxospore Ascorbic acid Bruckmayer-

nummuloides Berkenbusch 1954

Rhodophyceae
Porphyra sp. Asexual Purines Teramoto and

reproduction Kinoshita 1960


Porphyra Conchosporangia Plant growth Song 1991

yezoensis substances

Phaeophyceae
Saccharina japonica Zoosporangial Abscisic acid Nimura and

(=L. japonica) sori Mizuta 2002


Laminariales Zoosporangial IAA Kai et al. 2006

members sori




Chrysophyceae
Dinobryon cylindricum Encystment Vitamin B 1 Sandgren 1981


Biotic factors

Extracellular organic substances of algae helpful to reproduction

In some blue-green algae like Anabaena circinalis, A. torulosa, Cylindrospermum licheniforme, Nodularia spumigena and Westiellopsis prolifica, culture filtrates from sporulating cultures hasten the sporulation in the vegetative cells of the same alga (Table 9). But such was not the case in Anabaena cylindrica (Whitton 1965), Aphanizomenon flos-aquae (Rother and Fay 1979), Nostoc ellipsosporum (Ahluwalia and Kumar 1983) and Nostoc PCC 7524 (Sutherland et al. 1979). In many green and brown algae, female gametes or eggs released volatile secretions (pheromones/ gamones) which induced synchronous spermatozoid release and chemotaxis (Table 9).


Table 9 Alga may release some extracellular substance (s) in the medium which induced reproduction in the same alga/ other algae
________________________________________________________________________
Alga or Structure/ Induction helped by Reference

flora process induced



Cyanophyceae
Anabaena Akinete Culture filtrate of the Fay et al. 1984

circinalis alga
Anabaena Akinete Culture filtrate of the Fernandes and

torulosa alga (5- 8 d old) Thomas 1982
Cylindrospermum Akinete A chemical substance Fischer and Wolk

licheniforme (C7H5OSN) purified 1976; Hirosawa

from the centrifugal and Wolk 1979

supernatant fluid of a, b; Lang 1980;

sporulating cultures Van De Water and

Simon 1982
Nodularia Akinete Culture filtrate from Pandey and

spumigena sporulating culture Talpasayi 1980

of the alga


Westiellopsis Akinete Filtrate from 60- d Agrawal and

prolifica old culture of the Sharma 1996 b

alga



Chlorophyceae
Chlamydomonas Chemoattraction Pheromone (lurlenol) Starr et al. 1995;

allensworthii of male gametes secreted by female Jaenicke and

by female strains Starr 1996

gametes
Chlamydomonas Agglutination Gamone (glycoprotein) Moewus 1933;

eugametos of gametes liberated by female Förster et al.

gametes 1956; Lens et al.

1980
Chlamydomonas Mating Gamone produced by Tsubo 1961;

moewusii female gametes McLean et al.

1974
Chlamydomonas Sperm Extracellular sperm- Pascher 1931



paupera attraction attracting substance

of the alga


Chlamydomonas Zoospore An extracellular Mihara and

reinhardtii liberation substance of the alga Hase 1975
Cell wall An extracellular lytic Claes 1971;

lysis of factor from gametes, Adair 1985;

gametes, agglutinins (extrinsic Asamizu et al.

agglutination glycoproteins rich in 2000; Shrager

hydroxyproline and et al. 2003

serine)
Closterium Diffentiation Sexual pheromone Kato and Sasaki



ehrenbergii, of gametangial secreted by mtcells 1985; Fukumoto

C. peracerosum- cells, chemotactic et al. 1998;

Strigosum activity, migration Sekimoto 2000;

littorale of mt+ cells Tsuchikane et al.

towards mt- cells, 2003; Akatsuka

sexual fusion et al. 2006;

Yuki et al. 2010


Cosmarium Movement of Sexually- attractive Brandham 1967

botrytis cells for substance secreted

conjugation by cells


Dasycladus Mating Secretion of sexual Jollos 1926

clavaeformis activity substance by alga
Eudorina Sperm A dialyzable substance Szostak et al. 1973

elegans packet produced by male clone

Gayralia Gamete Substance extracted Moewus 1940 a

oxysperma formation from fertile sexual alga

(=Monostroma



wittrockii)
Oedogonium spp. Attraction of A species specific Tiffany 1957;

sperm and diffusible substance Hoffman 1960;

androspore secreted by oogonia Rawitscher- Kunkel

to oogonia or oogonial mother cell and Machlis 1962;

(named circein in Machlis 1962;

Oe. donnelli) Machlis and

Rawitscher- Kunkel

1967; Machlis et al.

1974; Hill 1980;

Hill et al. 1989;

Maier 1993



Protosiphon Sexual A substance secreted Moewus 1933

botryoides reproduction by alga
Sphaeroplea sp. Sperm Liberation of sexual Pascher 1931

attraction substance by eggs


Stephanosphaera Reproduction Sexual substance Moewus 1933

pluvialis secreted by alga
Tetraspora Reproduction Secretion of sexual Geitler 1931

lubrica substances by both

male and female

gametes
Volvox spp. Development Culture filtrates from Darden 1966, 1970,

of sexual sexual population 1971, 1973, 1974;

coenobia having species- Kochert 1968;

specific pheromones Starr 1968, 1969,

1970 a, b, 1971,

1972; McCracken

1970; McCracken

and Starr 1970;

Van De Berg and

Starr 1971; Pall

1973; Starr and

Jaenicke 1974;

Karn et al. 1974;

Kochert and Yates

1974; Meredith and

Starr 1975; Starr

et al. 1980;

Weisshaar et al.

1984; Wenzl and

Sumper 1986;

Balshüsemann and

Jaenicke 1990;

Hallmann et al. 1998;

Hallmann 2003




Xanthophyceae
Botrydium Sexual Sexual substance Moewus 1940 b

granulatum reproduction secreted by alga

Phaeophyceae
Ascophyllum Attraction of An attractant Müller et al. 1982 b

nodosum spermatozoids (finávarrene)

to eggs



Cutleria Attraction of Sex pheromone Derenbach et al. 1980

multifida male gametes (multifidene)

to source

release of

pheromone


Desmarestia Attraction of Sex pheromone Müller et al. 1982 c;

aculeata, male gametes (desmarestene) Maier and Müller

D. viridis to source 1986

release of

pheromone
Dictyota Attraction of Sex pheromone Müller et al. 1981

dichotoma spermatozoids (dictyopterene c’)

to eggs secreted by eggs


Ectocarpus Attraction of A volatile substance Müller 1967, 1968;

spp. male gametes (ectocarpene) secreted Müller et al. 1971;

to female by female gametes Pohnert and Boland

gametes 2002


Fucus spp. Chemotaxis An attracting substance Cook and Elvidge

of sperms (fucoserratene) 1951; Raper 1952;

secreted Müller and Jaenicke

by eggs 1973; Müller and

Seferiadis 1977
Laminariales Sperm release, Lamoxirene (cis- 2- Maier 1982;

(Laminariaceae, chemotactic cyclohepta-2´,5´- Maier and Müller



Alariaceae, orientation of dienyl-3-vinyloxirane) 1986, 1990;

Lessoniaceae) sperms towards a sperm-releasing Maier et al. 1987,

Examples: luring eggs pheromone 1988, 2001



Alaria, Chorda, (no species specificity

Eisenia, in the stereochemistry

Saccharina of the pheromone)

(=Laminaria),



Macrocystis,

Nereocystis,

Undaria,

Saccorhiza

Perithalia Sperm release A low-molar mass Müller et al. 1985

caudata and attraction pheromone secreted by

eggs
Syringoderma Male Male- attractant (viridiene) Müller et al. 1982 a



phinneyi attraction secreted by female gametes

Chrysophyceae
Dinobryon Differentiation A secretion by female Sandgren 1981

cylindricum of male gametes culture

Osmotic stress

Many algae formed physiological resting stages, resting cells, palmelloid condition, aplanospore, akinete, cyst or thick- walled cyst when subjected to desiccation (physical water stress) or increased osmolarity (physiological water stress, Table 10).

Zoospore (or gamete) formation in algae required presence of liquid water or liquid medium (Table 11) and the process was inhibited when algae were exposed to solid substratum or drought.


Table 10 Physical water stress (desiccation) or physiological water stress (increased salinity) induced the formation of aplanospore, akinete or cyst in many algae

Alga or flora Structure/ Type of water Reference

process induced stess

Cyanophyceae

Chroococcidiopsis Resting forms Desiccation Caiola et al. 1996

sp. of isolated cells




Nodularia Akinete Increse in salinity Jones et al. 1994

spumigena of growth medium

(obtained from

a lagoon)

Chlorophyceae
Boergesenia Aplanospore Concentrated Enomoto and

forbesii sea water (× 2.0) Hirose 1972
Chlorococcum Aplanospore Drought Bold 1930

infusionum
Chlorophyceae Thick wall Desiccation Coleman 1983

members (many) resting cells


Dunaliella Resting cyst High salinity Borowitska 1981;

salina Brock 1986;

Borowitzka and

Huisman 1993


Eudorina sp. Palmelloid Progressive Schreiber 1925

condition desiccation


Gonium sp. Palmelloid Progressive Schreiber 1925

condition desiccation


Haematococcus Akinete Desiccation Bold and Wynne

sp. 1978
Protosiphon Coenocyst Drying of Oltmanns 1922;



botryoides substratum Moewus 1933,

(intense sunlight, 1935

increasing

temperature)


Spirogyra Parthenospore Strong solution Klebs 1896

varians of sugar

Zygnema Thick- walled Slow desiccation West and

ericetorum cyst Starkey 1915
Zygogonium Akinete Very dry habitats Fritsch 1916;

ericetorum Hodgetts 1918

Xanthophyceae
Botrydium sp. Aplanospore Damp air Fritsch 1935
Vaucheria Aplanospore, Dry condition, Stahl 1879;

geminata, cyst drought Klebs 1896;

V. hamata, Puymaly 1922;

V. repens, Couch 1932

Vaucheria sp.

Bacillariophyceae
Freshwater Cyst Desiccation Round et al. 1990

Diatoms


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