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STUDIES ON SPORULATION AND PROPAGATION
IN SELECTED AGAROPHYTES
THESIS SUBMITTED
IN PARTIAL FULFILMENT OF THE REQUIREMENTSFOR THE DEGREE OF
DOCTOR OF PHILOSOPHYOF THE
UNIVERSITY OF COCHIN
BY
SHOBHA PRATAPRAO SHERE, M. SC.
CENTRE OF ADVANCED STUDIES IN MARICULTURE
CENTRAL MARINE FISHERIES RESEARCH INSTITUTE, COCHIN-682018
MARCH, 1985
CERTIFICATE
This is to certify that the thesis entitled ”STUDIESON SPORULATION AND PROPAGATION OF SELECTEDAGAROPHYTES” is the bonafide record of the workcarried out by Kumari SHOBHA PRATAPRAO SHEREunder my guidance and supervision and that no partthereof has been presented for any other Degree.
Dr. P. V. Ramachandran Nair, M.Sc., Ph.D.Scientist S-3
Cochin-682 018 Central Marine Fisheries Research InstituteMa rch-1985 Cochin-682 O18
DECLA RA Tl0.V
I hereby declare that this thesis entitled “STUDIES ONSPORULATION AND PROPAGATION IN SELECTEDAGAROPHYTES” has not previously formed the basis of theaward of any degree, diploma, associateship, fellowship or other
similar titles or recognition.
Cochin-682 018March, 1985. Shobha P. Shere
PREFACE
After passing M.Sc. in.Botany with Palynology as aspecial subject, the candidate joined the Central MarineFisheries Research Institute for the Ph.D. programme ofthe Centre of Advanced Studies in Mariculture in 1980.During the first semester she got familiarised with thevarious disciplines connected with mariculture such asibasic science, finfish culture, mussel culture, shrimpculture, seaweed culture, environmental monitoring andconnected field work and analytical techniques. Duringthis period the candidate was assigned the work on sporugla—tion and propagation in some agarophytes. After passing thequalifying examination with mariculture and algology as‘special subjects, the candidate moved over to the RegionalCentre of CMFRI, Mandapam Camp for collecting field data
and carry out the laboratory work.
During the stay at Mandapam the candidate had beenvisiting Kilakkarai, Thonithurai and Puthumadam fortnightlyto collect field data on four species of agarophytes viz.,Gelidiella‘ggg£g§§ (Forsskal) Feldmann.§tIHamel, Gracilarggcorticata J. Agardh,lQ. edulis (Gmelin) Silva, and Hxpneamusciformis (Wulfen) Lamouroux. In addition, the candidate
also visited the coral islands near Rameswaram Island in theGulf of Mannar where there are luxuriant algal flora.Because of its unique position with the Gulf of Mannerhaving a typical oceanic environment and ecologicalconditions on one side and Palk strait with eutrophicconditions on the other side, and the changing pattern ofthe monsoon and winds make one area or the other at Mandapam
always approachable for field work and sample collections.
The samples were regularly brought to the laboratoryand sporulation studies under different environmental
conditions were carried out in the laboratory for_§g¥§p;rexperiments.
The theme of the work is focussed on the spore outputstudies of some economic agar and Carragheenan producing redalgal species of Gelidiales and Gipartinales of Floridiophycae;
Gelidiella acerosa (Forsskal) FieldmannԤt_Hamel, Gracilariacorticata J. Agardh., Gracilaria edulis (Gmel) Silva; andHypnea musciformis (Wulfen) Lamouroux. The study was carriedout for two years from October 1981 to October 1983. Datawere collected on the seasonal and diurnal aspects of sporeliberation of the four red algae to know the spore liberationof the four red algae and to know the spore production potentialof the fertile fronds during different months of the year.
Experiments were also conducted in the laboratory tounderstand the effects of different environmental factors
such as desiccation, salinity, temperature, light-intensityand photoperiod on spore shedding. During these experimentssome information on spore germination has also been collected.
V11
ACKNOWLEDGEMENT
I wish to express my deep sense of gratitude toDr. P.V. Ramachandran Nair, Scientist S ~ 3 and Head,Fishery Environment and Management Division, CentralMarine Fisheries Research Institute for encouragementand guidance given during the research work, which enabledthis programme of work to materialise. I wish to expressmy most sincere gratitude to Dr. E.G. Silas, §ub-ProjectCoordinator and Director, Central Marine Fisheries ResearchInstitute, Cochin-18 whose suggestions and directions givenat different stages of the research study were equallybeneficial. The candidate extends her heart-felt thanks toDr. E.G. Silas for the same and for providing variousfacilities of the C.M.F.R. Institute, Cochin and the RegionalStation at Mandapam Camp till the completion of the thesis.
I offer my sincere thanks to Shri S. Mahadevan,Officer—in~Charge, Mandapam Camp and to the then.Joint
Director, Dr. P.S.B.R. James for providing necessary help
and various facilities during the research work at MandapamCamp. To Dr. N. Kaliaperumal, Scientist S - 2, who hasrendered advice, guidance, help and most of the supervisionat various stages of the research work during the research L’study at Mandapam Camp, I offer my most sincere gratitude.I also offer my sincere thanks to Dr. V.S.K. Chennubhotla,
viii
Scientist S - 2, for providing necessary help and guidanceand my thanks are also due to Shri Krishnapillai ShriS.Kalimuthu, Shri J.R. Ramalingam, Shri S. Selvaraj, ShriM. Najmuddin and other scientists and the staff of MandapamRegional Centre for extending help in various ways duringthe research work.
I wish to express my sincere gratitude toDr. P. Vedavyasa Rao, Scientist S - 3, Central MarineFisheries Research Institute, Cochin—18, for various suggestionsand encouragement during the course work of the programme andin the later stages of the research study.
I am especially grateful to Dr. C.V. Kurian, EmeritusScientist, Central Marine Fisheries Research Institute forgoing through the manuscript critically and offering manyhelpful suggestions at very short notice.
I express my gratitude to the ICAR/UNDP, New Delhi
for offering the Senior Research Fellowship to carryout theResearch work at the Centre of Advanced Studies in.Mariculture,Central Marine Fisheries-Research Institute, Cochin~18.
ix
§$gDIES ON SPORULATION AND PROPAGATION OF SELECTED AGAROPHYTES
§.9.flI§§..'-‘2.§
E252
DECLARATION iiCERTIFICATE iiiPREFACE ivACKNOWLEDGEMENTS vii
CHAPTER 1 INTRODUCTION . . . . . . . . . . 1CHAPTER 2 REVIEW OF LITERATURE . . . . . . 6CHAPTER 3 MATERIALS AND METHODS . . . . . 29
3.1 Spore Output . . . . . . . . . . 303.2 Spore Count , . . . . . . . . . 313.3 Seasonal Changes in Spore Output 333.4 Diurnal Periodicity in Spore Output 333.5 Effects of Environmental Factorson Spore Output . . . . . . . . 343.6 Germination of Spores . . . . . . 373.7 Seasonal Changes in the selectedEnvironmental Parameters . . . . . 38
CI-IAPTER4 R.ESULTS..........._..... 394.1.1 Fruiting Behaviour . . . . . . . . 394.1.2 Morphological appearance of the
fruiting fronds of the species ofO 0 0 0 o O 0 0 0 0 D 0 O l 0#02 SPOTE‘ 0 o o o o 0 0 an o o 9 o4.3 Seasonal Changes in Spore Output . . 44
4.4
4.5
4.5.14.5.24.5.34.5.44.5.54.6
4.6.1
4.6.2
4.6.3
4.7
CHAPTER.5.
CHAPTER 6.
CHAPTER 7.
Diurnal Periodicity in theSpore Shedding . . . . . . . . . . . 46Effects of Selected EnvironmentalFactors on Spore Shedding . . . . .. 48Exposure to Air and Desiccation . , 48S0.=lin_'L‘l:y..............Light Intensity . . . . . . . . . .. 51Temperature oouocooéolooPhO'tOperiOd 69000-9040000Germination of Spores . . . . . . . 55Seasonal Variations in SporeGermination oooooboooo-onEffects of Environmental Factorson Spore Germination . . . . . . . .Effect of Exposure to Air andDesiccation . . . . . . . . . . . .Seasonal variations in the hydrologicalparameters at the three collectionlocalities and their relationship"t0 SpOr'U.].atiCn¢o6ooooo¢ooDISCUSSION . . . . . . . . . . . . 69smmmny . . .. . .. . ..i. .. . 123REFERENCES.-............. ‘I23
Plate Nos.
I (A-D)
II A(B-C)
III AB
IV’ AB
C
D
V AB
VI A
VII
VIII A
IXX A
BCD
LIST OF PLATES
Thonithurai coast showing boulders andplatforms of compressed sand stones withseaweed growth.
Gelidiella acerosa, habit.Tetrasporio thalli.Q..ggg£g§§ with stichidia.Graqilaria corticata (tetrasporic) habit,pg. corticata (oystocarpic) habit.Q. corticata (tetrasporic), habit.Tetrasporio thalli of‘§. oortioata (magnified)Cystooarpic thalli of Q. gorticata.Cystooarpic thalli of'§. cortipata showingcystooarps.Tetrasporic plant of G edulis, habit.Cystocarpic plant of.§. edulis, habit.Tetrasporic plant of Q, edulis.C stocarpoc thalli of G. edulis. _ _ _ystocarpic thalli ofj§.'eEUIIB showing cystooarps.Hxpnea musoiformis, habit.Tetrasporic plant of HZpnea_ggsciformis.Cystocarpic plant of Hxpnea muspiformis showingcystocarps.Experimental sets kept for sporulation studies.Tetraspores of Gelidiella aoerosa.Spores of Gracilaria oortioatg.Spores of Eraciiaria edulis.Spores of Hzpnea musciformis.
1.
2.E
2.F
2.G
2.H
3.
4.
5.
xii
LIST OF FIGURES
Coastline of Mandapam showing the three collectionlocalities namely Thonithurai, Pudumadam and Kilakkarai.Seasonal variation in stichidia (A) and_tetraspores (B)of Gelidiella acerosa and tetraspores (c andcarpospores (D o racilaria corticata.‘Fruitinglfrond of GeIigiei1a'acerosa,tshowingietetrasporangiag stfafiidia.Cystocarpic frond of Gracilaria_9orticata showing thearrangement of oystocarps;s 'T*,eb:1;;:: ,_;,Cystooarpic_frond~of.g}U§dUIis snowdng*thnrarrangementof cystocarps.JEart of a tetrasporic frond ofiH nea musciformisto show_morphologicol,appedrance of EotraspofinngialStichidiaa
Rhythmic liberation of carpospores in Gracilarigmicorticata Ag G. edulis B and H nea musci?o1__§rom umadam'(C) and Kilakkarai (E).
Seasonal variation in the liberation of tetraspores (A)and carpospores (B) of Gracilaria edulis andtetraspores (C) and carpospores (D) of Hypneamusciformis.
Diurnal periodicity in the shedding of tetraspores of(Gelidiella acerosa collected from October 1981 toSeptember 19 .Diurnal periodicity in the shedding of tetraspores ofGelidiella acerosa collected from October 1982 toSeptember 1§§3.
Seasonal variation in the diurnal periodicity in theliberation of tetraspores of Gracilaria corticatacollected from October 1981 to September, 982.Seasonal variation in the diurnal periodicity_in theliberation of tetraspores of Gracilaria corticata‘collected from October 1982 to September) -.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
xiii
Seasonal changes in the diurnal periodicity in theliberation of tetraspores of Gracilaria eduliscollected from October 1981 to September 1985.Seasonal changes in the diurnal periodicity in theliberation of tetraspores of Gracilaria eduliscollected from October 1982 to September 1983.Seasonal variation in the diurnal periodicity in theshedding of tetraspores of H nea'pusciformiscollected from October 1981 o eptember 1985.Seasonal variation in the diurnal periodicity in theshedding of tetraspores of H nea musciformiscollected from October 1982 ¥o §eptember—19§3.
Diurnal periodicity in the carpospores output ofGracilaggg corticata collected from October 1981 to§eptember 1982.
Diurnal periodicity in carpospore output of Gracilariacorticata collected from October 1981 to September 1982.Diurnal periodicity in the shedding of carposporeof Gracilarig edulis collected from October 1981 toSeptember 1982.
Diurnal periodicity in the shedding of carposporesof Gracilaria edulis collected from October 1982 toSeptember 1983.
Diurnal periodicity in the liberation of carposporesof H nea musciformis collected from August 1982 toJuly 983.Effect of desiccation on the tetraspore output inGelidiella acerosa.Effect of desiccation on the liberation of tetrasporesand carpospores in Gracilaria corticata.Effect of desiccation on the liberation of tetrasporesand carpospores in Gracilaria edulis.Effect of desiccation on the shedding of tetrasporesand carpospores in.H1pnea musciformis.
22.
23.
24.
25.
26.
284
29.
50.
31.
32.
33.
34.
35.
36.
37.
Effect of salinity on the tetraspore output inGelidiella acerosa¢
Effect of salinity on the tetraspore and carposporeliberation in Gracilaria corticata;Effect of salinity on tetraspore and carposporeshedding in Gracilaria edulis.Effect of salinity on the discharge of tetrasporeand carpospore in Hxpnea musciformis.Effect of light intensity on the shedding oftetraspores in Gelidiella acérosa.Effect of light intensity on the tetraspore andcarpospore output of Gracilaria corticata.Effect of light intensity on the liberation oftetraspores and carpospores in Gracilaria edulis.Effect of light intensity on the tetraspore andcarpospore output of Hzpgea musciformis.Effect of temperature on the release of tetrasporesin Gelidiella acerosa.Effect of temperature on tetraspore and oarposporeshedding in Gracilaria_g9rticata.Effect of temperature on liberation of tetrasporeand carpospores in.Qg@cilaria edulis.Effect of temperature on tetraspore and carposporeoutput in Hgpnea musoiformis.Combined effects of photoperiod and light intensityon the tetraspore discharge in Gelidiella acerosa.Combined effects of photoperiod and light intensityon the tetraspore discharge in Gracilaria corticata.Combined effects of photoperiod and light intensityon carpospore output in Graci1aria_§grticata.Combined effects of photoperiod and light intensityon the carpospore output in Gracilaria edulis.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
Combined effects of photoperiod and light intensityon the tetraspore output in Gracilaria edulis.Combined effects of photoperiod and light intensityon the carpospore output in Hxgnea musciformis.Combined effects of photoperiod and light intensityon the tetraspore output in.HxEnea musciformis.Seasonal variations in the germination of tetrasporesof (1) Gelidiella acerosa and (23 Gracilaria corticata.Seasonal variations in the gerination of(1) carpospores of_Qraci1aria corticata (2) tetrasporeof Gracilarig edulis.Monthly variationsnin the germination of (1) carposporeof gracilaria_ggg;;§, (2) tetraspores oflgxggggmusgiigrmis. 'Monthly variations in the germination of carposporesof Exgggg musciformis (i) from Kilakkarai and(if) from Pudumadam.
,Effect of desiccation on spore germination :(A) Germination of tetraspores in Gelidiella acerosa.(B) Germination of tetraspore and carpospore in
Gfag1lartg*edgg; s(C) Germination of tetraspore and carpospore in
GHacilarié1edg;;$s.
Effect of desiccation on the germination of spores(tetraspores and carpospores) in Hzgneagmusciformis.
Effect of salinity on (1% germination of tetrasporein.§g;;g;g;;§ acerosa 2 germination nf tetrasporein Gracilaria corticata(3ggermination of tatbosporein'GracTIaria corticata 4 germination of tetrasporein_Gracilaria e u is.
48.
49.
50.
51.
52.
xvi
Effect of salinity on the following :(1) Germination of garpospore in Gracilaria edulis.(2) Germination of tetraspore in.H1Qnea musoiformis.
(3) Germination of carpospore in Hygnea musciformgg.Effect of temperature on the :(1) Germination of tetraspores in Gelidiella acerosaq(2) Germination of tetraspores(3) Germination of carpospores(4) Germination
in Gracilaria_g9rticata.in Gracilaria cortioata.
of tetraspores in Gracilaria edulis.Effect of temperature on the following :(1) Germination(2) Germination(3) Germination
Effect of light(1) Germination
(2) Germination(3) Germination(4) Germination
Effect of light(1) Germination
(2) Germination
(3) Germination
of carpospores in Gracilaria edulis.of tetraspores in.HxEnea)musciformis.of carpospores in H1Qnea_mpsciformi§.intensity on the following :of tetraspores in Gelidiella acerosa.of tetraspores in Graoilaria corticata.of carpospores in Graoilaria corticata.of tetraspores in Graoilaria edulis.intensity on the following :of oarpospore in Gracilaria edulis.of tetraspore in _};I1_p_r_;_e_a musciformi_s_.
of carpospore in.HZEnea musciformis.
53.
54.
55.
56.
57.
xvii
mbined ffects of light intensity and photoperiodLD cycle? on the following :
(1) Germination of tetraspore in Gelidiella acerosa.(2)“Sermination of tetraspore in Gracilaria corticata.(3) Germination of carpospore in Gracilaria corticata.Combined effects of light intensity and LD cycle on :(1) Germination of tetraspore in Gracilaria edulis.(2) Germination of carpospore in Gracilaria edulis.(3) Germination of tetraspore in Hxpnea musciformis.(4) Germination of carpospore in.H2pnea musciformis.Seasonal variations in the selected environmentalparameters at the 3 localities of collection.(e.g. Atmospheric temperature, surface.seawatertemperature, salinity, dissolved oxygen, and pHof the seawater in the collection localities)
at Thonithurai (Station I).Seasonal variations in the environmental parametersat Station II. (Pudumadam)Seasonal variation in the environmental parametersat Station III. (Kilakkarai)
xviii
LIST OF TABL§§
Tetraspore output on different days in Gelidiellaacerosa collected from Kilakkarai.
Liberation of tetraspores on different days inGracilaria_9orticat§ collected from Pudumadam.Tetraspore liberation on different days in Gracilariaedulis collected from Thonithurai.Tetraspore output on different days in Hxpneamusciformis collected from Pudumadam.
Shedding of carpospores on different days inGracilaria gorticata.Carpospore output on different days in Gracilariaedulis collected from Thonithurai;
Carpospore shedding on different days in Hxpneamusciformis collected from Pudumadam.
Carpospore output on different days in Hxpneamusciformis collected from Kilakkarai.Maximum and minimum values of spore output :obtained during the present investigation.Seasonality of spore production in differentmarine seaweeds.
CHAPTER - 1
INTRODUCTION :
The decade of 1970s heralded a major research effort intomariculture of economically important red and brown sea-weeds,particularly the agarophytes and alginophytes. The limitedand declining natural populations, lack of syntheticsubstitutes for the phycocolloids with their upwardlyspiralling costs have emphasised the need for more research
in this area. Now, algal mariculture has hecome technicallyand scientifically more feasible}
The marine red algal species (agarophytesj of Florideophyceae,(Rhodophyta) especially belonging to the orders Gelidialesand Gigartinales are used for the manufacture of agar-agar,carrageenan and other phycocolloids in many countries.“Agarophyte" is a term applied to those sea—weeds producingamorphous gelatinous substance referred as ‘agar’. InIndia Gelidiella acerosa and Gracilaria edulis are_used as
raw material for the manufacture of agareagar by thesea-weed industries. Apart from this, several genera ofthe order Gelidiales such as gpanthopeltis, Gelidiella,Gelidigm,‘Eterogladia and Suhria and those of the orderGigartinales like A ardhiella, Chondrus, Eucheuma, Gi artina,Gracilaria, Gygnogongrus, Hzpnea and Sarcodig are consumedas food either as raw or in processed form in different
countries particularly in the Indo-Pacific region. Thisobvious use of seawweed for food has reached its peak inJapan where different species are harvested for this purpose.
In Wales and Ireland laver (ggxgpg umbilicargis) is atraditional delicacy. In many countries seaweed biomassis used as animal fodder, fertilizer or as an energy sourceto produce methane gas. Seaweed extracts and composts areapplied to crops as a nutrient source andtaoil conditioner.(Subba Rao, 1965; Levring_gt_g;., 1969; Santelices, 197A;Michanek, 1975 and Chapman and Chapman, 1980).
Many genera and species of the orders Gelidiales andGigartinales are widely distributed in temperate andtropical to sub-tropical waters of the world. In India 6species of Gelidiella (Sreenivasa Rao, 1970 and SreenivasaRao and Trivedi, 1974), 17 species and 2 varieties ofGracilaria (Umamaheswara Rao, 1972 a) and 9 species of Hzpnea
(Untawale.g3.g;., 1983) have been reported. Many of thesespecies form an important component of the seaweed vegetation
at various localities along the Indian shores. Along thecoasts of Mandapam Gelidiella acerosa, Gracilaria arcuatavar. arcuata,;§. arcuata var._attenuata,;§. canaliculata,‘Q. corticatg var. corticata,.§..corticatg var. c lindrica,Q. cgvlifindrica, Q. disticha, Q. edulis,,'_(§. foliifera, Q,'b 9.! meinnarensistr 92 ! 9- .9.P...t.‘:.1.§.§o 9.0 t
IQ. verrucqsa, Hypnea musciformis,Ԥ. pannosa,Ԥ. spinella
and.fi. valentiae have been recorded by some workers(Umamaheswara Rao, 1969 and 1972 and Rama Rao and
Subbaramaiah, 1980).
A vast amount of literature is available on the morphology,
taxonomy, growth, reproduction and phycocolloid content09 some economic seaweeds e.g., Gelidiella acerosa of the
order‘§elidiale§ and various Gracilaria and Hypnea speciesof the Gigartinales order (Floridiophyceae, Rodhophyta) byFritsch (1945) Umamaheswara Rao, (1969 and 1972), Rama Rao
and Subbaramaiah, (1980), Rao, (1970), Rao and Krishnamurthy(1968), K. Rama Rao (1970 - 1982). But the informationsthat are available on spore producing capacity of the algaeare scanty particularly with reference to the effect ofenvironmental factors on spore shedding and the germinationof spores. In order to assess the mariculture potentialof these species by spore mihod, sporulation studies on thesame were undertaken on_Qelidiella acerosa, GracilariaCOrt;9a§§, G._ggg;;§ and Hypnea musciformis growing alongI’.'."3
the Mandapam coast.
Studies on spore phenomena have been initiated in someeconomic algae such as Gelidiella, Gelidium, Porphyra,Gracilaria and Hypnea of Rhodophyta and Dictyota,
Turbinaria, Padina species of Phaeophyta and in'g;y§ speciesof Chlorophyta by Rao, (1971 a b), Tanaka, (1941), Mshigeni,(1976), Suto, (1950), Yinam Ngan and Price, (1983), Detailedknowledge on sporulation is not available as regards the othermembers of Gelidiales and Gigartinales occufiing along theIndian coasts.
During the present work, carried out for a period of 2 yearsfrom October 1981 to September 1983, data have been collectedon the fruiting behaviour and experiments were conducted atroom temperature on the daily and seasonal changes in the
shedding of tetraspores of Gelidiella aoerosa and tetrasporesand carpospores of Graoilarig corticata,.§. edulis andHypnea musgiformis and also periodicity in the daily liberationof spores in these four red algae, Influence of importantenvironmental factors such as desiccation, salinity, lightand temperature on the spore output have been studiedin det 0
Data have also been collected on germination of spores offour red algae. Informations on the hydrological conditionsof the inshore waters in the vicinity of Mandapam such assurface sea-water temperature, pH, salinity and dissolvedoxygen were also collected. Results obtained on theabove aspects are presented and discussed in this thesis.
CHAPTER - 2
R§yIEw OF LLIERATURE
The red algal members of Florideophyceae (Rhydophyta)
especially belonging to the orders Gelidiales andGigartinales ocouring in various regions of the world havebeen studied for the past four decades in view of their
commercial value as a sources of agar—agar and human food.These investigations carried out on different species of
Gelidium and Eterooladia (Santelices, 1974), Gracilaria(Hoyle, 1975) and.§zpggg (Mshigeni, 1976 a, b,_d and 1977and 1978) and Mshigeni and Lorri (1977) provide pertinent
information to our knowledge on their taxonomy, biologyand economic importance.
Studies have been conducted in India on some importantmembers of the order Gelidiales and Gigartinales. Amongthe order Gelidiales, studies were carried out on theecology, commercial utilisation and cultivation ofQelidiella acergsg, by Sreenivasa Rao, (1969, 1971 a, b, c,1974 and 1976); Umamaheswara Rao, (1973 a and 1974 a);
Sreenivasa Rao and Trivedi, (1974); Subbaramaiah §§‘g;.,
(1974 and 1975); Bhanderi, (1974); Krishnamurthy.g§_a;¢.((1975); Subbaramaiah.§§‘g1., (1975): Thomas g§'g;.;(1975
a and b); Rama Rao.g§.§l.,(1976); Chennubhotla §3_g;.,(1977a, b and 1979); Rama Rao and Subbaramaiah,(1977
and 1979); Joshi and Chauhan, (1979); Subba Rao, (1979);
Mairh_g§‘g;., (1979 a, 5); Pate1.g§'g;.,(1979 and 1980)other species of Gelidiella (Subba Rao, gt g;., (1977)and Untawale» .932 an (1977). 9232.92.92 .I.?.9.~“>.~°:...i..3.L.,_.u2 2.9.9
Pterooladia heteroplatos by (Umamaheswara Rao and
Sreeramulu, (1964) Mairh and Sreenivasa Rao, (1978);
Kaliaperumal, (1979); Kaliaperumal and Umamaheswara Rao,
(1981); and Umamaheswara Rao and Kaliaperumal, (1983).
Among the members of the order Gigartinales, investigations on taxonomy, morphology and anatomy were carriedout in certain species of_Gracilaria by Ahmed, (1966);
Oza and Krishnamurthy, (1967 and 1968); Krishnamurthy
g§‘§;., (1969); Raju and Thomas, (1971); Umamaheswara fiao,(1972 b, 1973 b, c, 1974 b and 1976); Rama Rao and Thomas,(1974); oza, (1975, 1976, 1978 and 1979); Subbarangaiah
_g§'g;., (1975); Thomas and Krishnamurthy, (1976); Thomas.(1977); Rama Rao, (1977 a); Mohan Joseph and Krishnamurthy,
(1977); Chennubhotla g§‘g;.,(1978 and 1979); Subba Rangaiah,(1978) and Umamaheswara Rao and Subba Rangaiah, (1980)
and.fi1pggg by Rama Rao (1970, 1972, 1976, 1977 a, b, 1979,and 1982); Rama Rao and Krishnamurthy, (1968 and 1978);
Subba Rangaiah (1978); Rama Rao and Subbaramaiah, (1980);
Umamaheswara Rao and Subba Rangaiah, (1980); Solimabi go a_l.,
(1980)., Rama Rao g_{; a_]; (1983),
2.1 SEASONAL GROWTH BEHAVIOUR
The plants of Gelidiella acerosa in the Palk Bay nearRameswaram and in the Gulf of Mannar near Pudumadam have
been observed throughout the year by Umamaheswara Rao,
(1973 a) and these plants attained maximum size in twoseasons with a half yearly growth cycle. The main growthseason with maximum number of large sized fronds was foundbetween December and April on the Palk Bay side and between
July and August on the Gulf of Mannar side by Umamaheswara
Rao, (1973 a). Gelidiella acerosa occurring on the reefat Kilakkarai reached maximum growth in November, as
measured by the length, bushiness and the number of branches
of plants (Thomas_g§_§;., 1975 a) and the plants showed2 peak growths for the same period, one in July and anotherin November, as measured by wet and dry method (Thomas.g§
g;., 1975 b). The peak values in monthly growth rates inlength and percentage cover of_§elidiella acerosa growingat Ervadi near Kilakkarai were recorded during summermonths May to July. Higher growth rate in cover was alsoobserved during winter months November to December (Rama
Rao and Subbaramaiah, 1979). Subba Rao, (1979) observed
that the mean monthly growth rates of the plants takenindividually or as in population were maximum in Januaryat Ervadi and in February at Rameswaram. Chennfibhotla
,g§_§;., (1979) also observed the vegetation of Gelidiellaacerosa occurring throughout the year at Pudumadam,Kilakkarai and Krusadai Island.
The seasonal growth behaviour of Gracilaria corticatagrowing in the Gulf of‘Mannar tide near Mandapam was
studied by Umamaheswara Rao (1972 b). He stated that theseplants occurred throughout the year with its peak growth
from.June to September and another small peak in Novemberor December. Similarly Chennubhotla.g31g;., (1979)reported the occurrence of Gracilarig corticata in allthe months of the year from Pudumadam;
Population of Gracilarig edulis in the Palk Bay nearRameswaram was also observed throughout the year by
Umamaheswara Rao (1973 b). Increase ingmean length was
observed by him from November or December and a large
-number of plants reached their maximum height during theperiod January to April, after which there was a declinein the mean length particularly during May and June.This reduction in the height seemed to be due to thebreakage or removal of fully grown and old fronds(defoliation) and_also by the development of fresh plantsin the population. Again another small or secondary peakin growth was found_in August and September with increasein height from July; During October/November there was
10
a considerable decrease in the average length of the plants.Similarly Chennubhotla‘gt_§;., (1979) also recorded thevegetation of Gracilaria edulis throughout the year atRameswaram and Krusadai Island.
2.2 FruitingBehaviour :Tetrasporophytes were observed only for some months in the
population of Gelidiellg acerosa growing at Veraval bySreenivasa Rao, (1974) and at Rameswaram by Umamaheswara
Rao (1973 a) and Rama Rao;gt.g;., (1976).
In Gracilaria corticata growing at Veraval, tetrasporicphase occurred almost throughout the year while sexualphase was seasonal.occurring from September to February(Oza, 1979). Tetrasporic and cystocarpic plants of
Gracilaria edulis occurred through the year in the Gulfof Mannar side at Mgndapam (Umamaheswara Rao, 1976) andVisakhapatnam coast (Subba Rangaiah, 1978). In Gracilariaedulis growing at Rameswaram in the Palk Bay tetrasporophytes occurred in all the months except in October orNovember. Antheridial and cystocarpic plants were seenonly in the month of January and only one or two plantswere observed in the samples examined (Umamaheswara Rao,1973 b).
11
2.3 SPORE PRODUCTLQN POTENTIAL IN ALGA
The spore production potential has been investigated in afew economic seaweeds, Suto (1950) developed a method forcounting the number of spores produced by seaweeds over agiven period of time.. In this method, a known size of asporulating thallus was collected from the field, broughtto the laboratory, and placed in a Container of seawaterwith a glass plate at the bottom. The spores liberatedfall on to the glass plates The plate containing the sporesis brought under the microscope and the spores are countedat hourly intervals for 12 hours. The mean value of thecount is then used for estimating the rate of spore kiberationand the total number of spores which a mature thallus canproduce. This method involves the counting by rows and hasbeen called (Katada 1955) the ‘obi’ method.
Suto (1950) in his counts on the spore production poten~tial in Gelidium amansii found that 1.0 g of a fertile
Afrond of this species produced about 10 to 105 sporesper day. Oza and Krishnamurthy (1967) using the samemethod (with slight modififiations) found that in.§§gg;lgg;§
ggggggggg a single mature oystocarpic plant produced up to1.97 x 104 carpespores per day in December, the season<fmaximum spore shedding. Similarly, Rao (1971 a) found thatan average seized tetrasporic thallus of Qelidiella acerpsgh
12
produced 2 x 104 spores per season (i.e. October — November).
Neushul (1972) developed a different method in estimating
the spore-production potential in.fl§crocystisi£g£ggg£§,by measuring the number of sporangia per square millimeter of the sporangial area and multiplying this by 32,the number of spores in each sporangium. He estimated
3.5 X 105 spores per mm2 of sporangial area on each sideof sporophyll (Anderson and North 1966).
In.§gggg£hi§§_germat9dea, individual sporangia were foundto contain 128 zoospores (Norton 1972), in Gracilariaverrucesg about 200 to 2000 carpospores can be obtained(Kin 1970) from a single cystocarp, while in.§§lggiggnebustgmfi 65 to 430 carpospores may be obtained (Guzman
del PPOO 2;. g;., 1972) from a single cystocarp and thetotal number of spores produced by a mature thallus may
be 3#,000 to nearly 300,000 carpospores. In most otherseaweeds like Gracilaria cortioa£§,‘g. ggggig and Hypneamusciformis occurring at Mandapam coast, this kind ofinvestigations has not been conducted.
The reproductive capacity of the two species of Hypneaspecially L §§Eusciformi_§ at Veraval, West coast oflndiaANDl§. valgntiae at Pamban and Krusadai Island, Eastcoast of India was studied over different seasons by Rama
13
Rao (1977 b). The tetrasporophytes in Hypnea muscifonggoccurred throughout the year with no distinct peak whilecystocarpic plants occufied only in October each year. Thetetrasporic plants in.Hypnea valentiae also occurred inall months at Pamban and Krusadai Island while the
cystocarpic plants occurred for 11 months at Pamban and for7 months in a year at Krusadai Island. Maximum frequency
of tetrasporophytes and carposporophytic plants of Hypnea_va1entiae at Pamban and Krusadai Island occurred in
February and June/July respectively. Male plants were notobserved both in Hypnea musoiformis andifi. valentiae. AtVisakhapatnam coast tetrasporophytes and carposporophytes
in the population of Hzpnea musciformis occurred in all themonths of the year (Subba Rangaiah, 1978).
2.4 SPORE SHEDDIQ: :
Information available on spore shedding in natural and indifferent environmental conditions influencing sporeshedding is very little on red algal genera. Japaneseworkers paid more attention to this aspect and collected
data onigglggigg, Porphyra and other economically importantseaweeds. The spore output estimated by Suto (1950 a and b)for the fronds of Gelidium varied from 1,00,000 to 10,00,000spores/g. fresh wt./day and the spore shedding was found
14
chiefly for one day, Boney (1960) estimated spore outputin Antithagnion plumula for a number of days with morethan 50 % spore release in the first 24_hrs, about 20 %on the second day and 16 % after 3 days.
Studies have been conducted on sporulation of some red
algae growing at Mandapam area and other localities ofIndian coast. Gelidium pusillum and Pterocladia heteroplatosat Visakhapatnam coast were studied by Kaliaperumal(1979).Gracilaria corticata from.Mandapam region was studied by
Umamaheswara Rao (1976) and.Mohan Joseph and Krishnamurthy(1977). Rama Rao and Thomas (1974) collected Gracilarigedulisxfrom Krusadai Island near Mandapam and studied
carpospore output. Krishnamurthy (1967) reported tetrasporeoutput in_Graci1aria millardetii. Oza and Krishnamurthy(1968) observed liberation of carpospores per plant inCracilaria verrucosa growing at Kuda near Bhavnagar in
Gujarat. The tetraspore and carpospore shedding in Hypneavalentige from.Mandapam was studied by Rama Rao (1979),
Subba Rangaiah (1978) collected data on tetraspore and
carpospore output in four members of Gigartinales order.namely Gracilaria corticata,Ԥ. textorii, Gracilariopsisgjoestedtii and Hxpnea musciformis growing at Visakhapatnamcoast. Shedding of tetraspores from stichidig in
15
Gelidiopsis.variabilis was observed for 3-4 days atVisakhapatnam coast by Kaliaperumal and Umamaheswara Rao,
(1982).
2.5 SEASONAL CHANGES IN SPORE OUTPUT :
Seaweeds vary tremendously in the timing of their sporeproduction. In some of the warm eastern shoreline oiSouth Africa (Isaac and Hewitt, 1953) or the tropicalregions such as India (Rao 1970) where the growth of gypggg,occurs throughout the year, ietrasporangia are borne onthe thalli throughout the year. Dictyota dichotoma andCentroceros plavulatum in India also produce spores(Umamaheswara Rao and Sreeramulu 1970) throughout the year.
In Gracilaria verrucosa, spore maturation was observed only(Oza and Krishnamurthy 1967; Ogata 1972) at certain periodsof the year. In this alga the cystocarps were found inSeptember or October in India and the peak spore productionwas attained_in December or January. Tetrasporic thalliin contrast was seen only from March to September. InGelidiella gggggga, there were observed (Rao, 1971 a)two spore-shedding seasons a year in India: (from Aprilto May and from October to November), each spore sheddingseason lasting 25 to 30 days. In Baja California, Gelidium.§gQg§tum has its peak for spore production during August
16
and September (Guzman — del - Proo.g§.g;., 1972), thetotal number of spores released gradually decrease towardsJanuary and February, coinciding with winter.- In manyalgae, the season of fertility is related to thegeographical location where the alga grows, and this maybe correlated with temperature or day length.
Seasonal changes in spore output in some seaweeds havebeen reported by few workers. Sreenivasa Rao (1971 a and1974) studied seasonal variation in tetraspore output inGelidiella acerggg growing in the tide pools at Veravalyg§;;g;gl;§.ggg£g§§ from Pudumadam was recorded byUmamaheswara Rao (1974). Kaliaperumal, (1979) collected
data on the seasonal spore shedding in Gelidium.3g§;;;ggand Pterogladia'he§g§gp;ggg§. Data on seasonal changesin tetraspore and carpospore output of Gracilariaoorticata growing at Mandapam were observed by Umamaheswara
Rao (1976) and.Mohan Joseph and Krishnamurthy (1977).
The seasonal change in tetraspore and carpospore output
in Gracilaria oorticata and gypnea musciformis growing atVisakhapatnam coast was also observed by Subba Rangaiah,(1978), Oza, (1979) conducted laboratory experiments inGracilarigdcorticata occurring at Veraval coast. Heobserved two peak periods of sporulation. Studies on the
17
seasonal rhythm in the shedding of tetraspores andcarpospores in Hypnea valentiae from.Mandapam werecarried out by Rama Rao (1979)¢= Maximum number of
tetraspores was observed in February and carpospores inOctober.
The spore shedding in Qracilaria edulis growing atRameshwaram was reported by Rama Rao and Thomas, (1974).
He stated that the total spore output per plant showedpeak values in July and August and gradual decrease byJanuary, He further stated that higher values of sporeoutput were seen again in FebruaryéMarch while inApril4May a total lack of spores was observed. The sporeshedding in Graoilaria vérrucosa was further observed by‘Jones (1959 a) and Oza and Krishnamurthy, (1968) andin Gelidigpsis variabilis by Kaliaperumal and UmamaheswaraRao, (1982),
18
2.6 DIURNAL PERIODICITY IN SPORE SHEDDING :
Diurnal periodicity in spore liberation was observed insome members of the order Gelidiales and other red algaeby few investigators. Suto (1950 b) and Katada.g:.g;.,(1953) observed diurnal shedding of spores in Gelidiugamansii daily in the afternoon.- Fukuhara (1957) conducted
diurnal experiments in.Iridophycus cornflcopiae and diurnalvariation in shedding of spores was scarcely'recognisedQMatsui (1969) observed diurnal periodicity in the liberationof tetraspores and carpospores in Gloiopeltisitgggg and inGloiopeltis furcata. In the former maximum release of sporeswas noticed from evening to midnight and in the latter inthe early morning. Umamaheswara Rao (197# a) reporteddiurnal periodicity in the liberation of tetraspores inGelidiella acerosa and in Gracilaria corticata (1976).Diurnal periodicity studies in the liberation of sporeswas also observed in Gelidium_pusil1um,_§e1idiopsis variabilis
and.in.§teroclag;g heteroplatos by'Kaliaperumal,-(l979)gA definite diurnal periodicity in the shedding of carposporesin_§ypgg§ valgntiae was reported by Rama Rao,*(1979);
19
Recently Ngan and Price (1983) collected data on the diurnalperiodicity of carpospores and tetraspores discharge in 14red algal taxa growing in the vicinity of Townsville region,Queensland, Australia under a variety of laboratoryconditions. Experiments were conducted at hourly orbi-hourly intervals over periods of 24 hours or in somecases 48 hours.
Differences in the time of peak shedding between tetraspores
and carpospores was observed by some workers. Katada.g§_a;.,(1953) in Gelidium amansii, Subba Rangaiah (1978) in Hypnea
musciformig, Matsui (1969) in Gloi0peltiS_£gg§§ aandgloiopeltis furcata; Umamaheswara Rao, (1976) and SubbaRangaiah, (1978) in_Gracilar;§ corticata, in Gracilariatextorii and Graoilariopsis sjoestedtii by Subba Rangaiah(1978) and in Gelidiumlpggillum and Eterocladia heterqplatosby Kaliaperumal, (1979).
;Seasonal variations in the diurnal periodicity of sporeoutput were investigated by Katada'gt_g;., (1953) inGelidium amansii. Similarly in Gelidium pgsillum andGelidiopsis yariabilis seasonal changes in the diurnalperiodicity of tetraspores was observed by Kaliaperumal(1979)oIn Gracilaria corticata,‘g. textorii, Gracilariqpsigsjoestedti; and Hypggg musgiformis the diurnal periodicity
20
of tetraspores and carpospores was reported by SubbaRangaiah, (1978). He observed that the diurnal periodicity_(La . ’ Vof tetrasports not vary in different months or seasons/\
of the year.
2.7 .E»\FFECTSW‘_C_)F’ENVIRO1\1I‘/IJ’:."N'Z[‘AL FACTORS ON SPORE SHEDDING :
Effects of environmentalfactors like desiccation, salinity,light (intensity and photoperiod) and temperature on sporeshedding have been further observed by some workers.
Since exposure to air and desiccation of plants caused by
tidal action influence spore production (Suto, 1956 a)experiments_were carried out by Katada (1955), Matsui (1969)and others, to study its effect. Katada (1955) reportedthat desiccation in the shade has no including effect upon
the shedding of tetraspores in Gelidium amansii and inthese experiments the time of shedding was extended withinabout halfea-day. On the contrary} Matsui (1969) reportedaccelerating effect in Gloiopeltis species.
Sreenivasa Rao, (1971 a) further reported that drying oftetrasporic plants in shade had no effect on spore outputin Q-elidiella acerosa, Umamaheswara Rao, (1976), (in.ggggiggggg‘gg£3igg£g)stated that tetraspore output decreasedmarkedly in thalli exposed for one hour and there was nospore shedding with fmrther increase of exposure to air.
21
While carpospores-were not at all liberated from cystocarpsduring one hour exposure at room temperature. RecentlySubba Rangaiah, 1978 and Uamaheswara Rao and Subba Rangaiah
(1980) observed the effect of exposure to air of thetetrasporic thalli in Gracilaria corticata, Gracilariaptextorri, Gracilariopsis sjoestedtii and Hypnea musciformis.The tetraspore output decreased in all plants with increasein the duration of exposure of the fronds and maximum sporerelease was observed under continuously submerged conditions.
Recently Umamaheswara Rao and Kaliaperumal, (1983) studied
exposure to air of tetrasporic fronds of Pterocladggheteroplatos, Gelidium pusillum and_gg;;diopsis variabilisgrowing at Visakhapatnam.coast; They observed maximumspores_in submerged conditions. Umamaheswara Rao and
Sanjseva Reddy (1982) studied tetraspore shedding of thetetrasporic fronds in Dictyota dishotoma with maximum
spore output in submerged conditions.
Yamasaki £3 gl., (1957) studied the influence of sea waterof various concentrations on the growth, formation ofsporangia and liberation of spores from the conchocelis
phase of_gg£pg1g§;t§g;g. He found more number of sporesin 28.19 %u3 whereas Matsui (1969) found that tetrasporeliberation was not significantly influenced in salinities
22
between 17 %o and 52 %o in.Q;piop@1ti§J§ggg; and Gloiopeltig
furcata.. Monospore release was found in Agrochaetigmggggphytioum by White and Boney (1969) in the media of
19.2 %o 9 49 %oS. Normal production of zoospores orgametes was found in _I=J__1_y_a_ §_a_§_9_igLa by Mohsen _e__1_:_ _a'_l_., (1972)
in the salinity range between 20 and 35 %o.. Higher salinitiesfrom 35 %o to 45 %oS enhanced both the formation of swarners
and their discharge.
The spore output experiments in different salinitiesranging from 10 ~ 60 %o were conducted by Subba Rangaiah
_e_1_;_ _a_Z_L_., (1975), Subba Rangaiah (1978) ‘and Umamaheswara Rao
and Subba Rangaiah (1980) in which the optimum range observedfor peak shedding of spores was 30 -"40 %o in Qracilaria
corticata,.g. textorii and Gracilariopsig_gjoestedtii andin.H1pnea.mg§g;;g§g;§ maximum spores were obtained at
20 - 30 %o. Umamaheswara Rao and Kaliaperumal (1983) alsoconducted spore output experiments with tetrasporic thalliof Gelidium pusillum, Bterocladia heteroplaggg andGelidiQpSiS gggggpgggg in different salinities rangingfrom O %o to 70 %o. In _(_I-__e_e_l_:i_._c_1_;i._}_;I_m and gterocladia
heteroplatos spore shedding was found at 10 %o — 60 %oand peak discharge at 30 %o salinity was obtained. InDictyotgigichotoma (Umamaheswara Rao and Sanjeeva Reddy£1982) maximum spore shedding was observed at 30 %o Salinity..
23
The effect of light intensity on spore shedding has beenstudied by some workers. White and Boney (1969) studied
the production of monospores in Acrochaetium endophyticum.
Massive spore output over long periods was seen in thelight intensity range of 50 - 110 lumens/Sq.ft.(538-1184 lux)in.yonostroma by Ohno (1972). The effect of light intensitywas further studied in.flpnostroqa nitidgg by Ohno andNozawa, (1972), in the tetraspore shedding of Gracilariacorticata, Gracilaria textorii and Gracilarigpggg sjoestedtiiby Subba Rangaiah (1978) and Umamaheswara Rao and Subba
Rangaiah (1980). In Graoilaria corticata,.g..§g§gQg;;.and Gracilariopsis sjoestedtgi maximum release oftetraspores were obtained in complete darkness, but inHyggea muscifogmig spore output was higher at 750.1 50 luxlight intensity than in total darkness. UmamaheswaraRao and Kaliaperumal (1983) found peak liberation of
tetraspores in Gelidium Eusillum, Pterocladia heteroplatos,and Gelidiopsis variabilgg at 500 lux.
The effect of day length on sporogenesis has been reviewedby Dixon and Richardson (1970) in.§ggg;g_fuscopurpures.
In this the conchicelis phase produced monosporangia onlyin short day.photoperiods. Iwasaki, (1961) observedmonospore formation and release of fertile EOHOSPOTQS in
24
Porphgra;§ggg;g, induced by short day conditions. Work onphotoperiod induction and liberation of monospores from
the conchocelis phase in 5 species of Porphyra was observedby Kurogi and Suto (1962): Porphyra tenera,l§. kuniedai,_]§_°_..p ‘ezoensis, §_. pseudolinearis and E. anggsta. Liberation
of monospores were also observed in short day conditions.Kurogi and Suto (1967) observed more liberation of monospores‘in Porphyra umbilicalis under long day conditions. InAcrochaetium endophyticum plants kept under long dayconditions produced large number of mcnospores at all lightintensities ranging from 30 — 500 lumens per sq.ft.(323 - 5382 lux). Ohno (1972) studied gamete liberationin.Monpstroma and found that long day treatment producedfaster gamete liberation than short day treatment.Umamaheswara Rao (1974 a) observed maximum shedding of
tetraspores at 16 +'B LD cycle in Gelidiella acerosa.Liberation of tetraspores and carpospores in Gracilaria
corticata was observed by gmagghggwgrg gag (1976). Hereported maximum tetraspore shedding in O + £1 £5 cycleand carpospore output in 4 + éaiffi cycle. Similarlymaximum release of tetraspores was observed by Subba
Rangaiah (1978) and Umamaheswara Rao and Subba Rangaiah
(1980) in Gracilariaflggrticata, Gracilaria textorii,Gracilariopsis sjoestedtii and Hypnea musciformis at O¥§H LD
25
cycle, and the spore shedding decreased with increase inthe duration of photoperiod. Recently Umamaheswara Raoand Kaliaperumal (1983) studied the combined effects oflight and its day length on the shedding of tetrasporesin .§.§.=‘.l.?l.*E1.?f.-..1.1...ml.P..1§..§>.5.L.3;.1._11.I.11: .E.:v.e-:r‘_o..<;1.-..€2_§1.i.s Bsteroplatos and
Gelidiopsis variabilig. Maximum spore output was obtained
at light intensity of 500 lux and photoperiod of 16 :'§and 2.4 :' '5 LD cycles.
The relation between spcrulation in seaweeds and temperatureof seawater was studied by Suto (1950 a and b). According to Suto (195C a) spore Shedding in seaweeds takesplace when the seawater temperature has reached an
optimum level for each species. He observed that sporeliberation in.§g;ggggg‘gggg§;;_takes place in the afternoon. Katada (1955) supported Suto's observation, alsoworking on Gelidiales for Gelidium.amansii. He statedthat shedding time of tetraspores_and carpospores variedaccording to the temperature of the seawater in the field.Fukuhara (1957) conducted experiments with Iridqphycus
cornucopiae. gurggg andiggiyamg (1966) examined the
effect of water temperature on the liberation of monospores on 6 species of Porphyra, namely §,‘§§gg£g, ‘Q.. 3. - . !. ~ » !. - ax;e.9;b;§.9l:t1,:e andE; _pseudolinearis. Monospores were liberated between 10°
26
and-25°C in.§, ggpera,'§.,yezoensis and P an usta, and0‘L8
liberation of spores was little or not seen.in.§..5gg$§dai,Q. suborbiculatg andl§.‘pseudolinearis at these temperature
ranges. Kurogi g§‘g;., (1967) noticed differences in theoptimum temperature of monosporexliberaiion between theconchocelis occurring in the autumn and spring plants ofPorphyra umbilicalis. Subba Rangaiah, (1978) andUmamaheswara Rao and Subba Rangaiah,(1986) observed
variation in the liberation or tetraspores at 5 diiteronttemperatures ranging from 15°C to 35°C in Gracilaria
corticata,_Gracilaria textorii and H1pnea‘mg§gi£ggg;§.Similarly the tetraspore output also varied at 9 differenttemperatures ranging from O'- 45°C in the tetrasporicthalli of Gelidium ousillum, Pteroqgggig heterqplatosand Gelidiopsis variabilis as observed by Umamaheswara Raoand Kaliaperumal, (1983). Peak output of spores wasnoticed at 250C in Gelidium pusillum and_§terocladiaheteroplatos and at 30°C in Gelidiopsis variabilis,Similarly Umamaheswara Rao and Sanjeeva Reddy, (1982)
observed maximum liberation of tetraspores from 25°C
to 309C in.Qggt1ota dichotoma.
27
2:8 GERMINATION7QF SPORES :
Some workers observed germination of spores in differentseaweeds. Barilotti and Silverthorne (1972) observedseasonal spore germination in Gelig;gg‘gQQg§3gQ from BajaCalifornia. Fruiting plants were seen throughout the yearand spores liberated during the spring and early summerwere germinated producing young thalli on new substrata.
In Chondrus crispus maximum germination rate was observed
as the early féllni éubba Rangaiah (1978) also observedmonthly variation in the number of dividing tetrasporesand carpospores in Gracilaria cortigata and Hypneamusciformis liberated on the first day and spore germinationwithin 24 hours from December 1974 to April 1976.Kaliaperumal, (1979) collected some preliminary data on the
percentage frequency of germinating tetraspores andcarpospores in Gelidium pusillum and tetraspores inPterocladia heteroplatgg. He did not observe dividingspores within 24 hours in Gelidiopsis_gggggg;;;§ duringthe two and a half years seasonal study on spore germination.
28
Spore germination have been shown to be influenced byenvironmental factors such as desiccation, salinity, lightand temperature by some workers. In Gelidium pysillugand _1_-;t_e_:;-_9_c1adia _1_1_eteroplat§§_ Kaliaperumal, (1979) found
maximum spore germination in controls than when exposed
to air. Subba Rangaiah, (1978) studied germination ofspores of Gracilarig corticata, and Hypnea musciformisin different salinities, Kaliaperumal (1979) foundmaximum germination of tetraspores of Gelidium pusillumin 20 %o and 30 %o whereas there was no germination in
the tetraspores of Pterocladia_heteroplato§é
The effect of light intensity on the germination ofcorpospores of Gracilari§‘y£§ggggg was studied byiJones (1959). The same study was also done by Whiteand Boney (1969( in_§g£gQaeg£gg endophyticgg, by
Subba Rangiah (1978) in _C;‘;_1;_;_g_;1____J1:_g_;_;;_L_§ _g_g_:;;t;.__%1;_g_ and
Hypngaimuscifgrmis, by Kaliaperumal (1979) om gglgdggg
'pg§;llgQ and Pteroclagig heteggplggg. Katada (1949and 1955) observed the effect of temperature on germination of carpospores and tetraspores in.§§;g§ggg amansii.Further studies on this aspect was done by Fukuhara,(1958) in Iridophycus corngggpigg, by Ohno, (1969) in.<2.9.1.i.di:Lg,m saarsii. by Subba Rangaiah, (1 978) in .§.2:.9£_i.3L.9~.;‘.i.a._
and gypgeg from Visakhapatnam coast, by Kaliaperumal,
(1979) in Qelidium_pusillEg and PtergQl§g;g_Qgfieroplatos.
29
CHAPTER -A3
MATERIAL AND METHODS
Mandapam (79O8'E, 9O17'N) is situated on the southern partof the East coast of India (Fig.1). To the north of Mandapamis Palk Bay and to the south the Gulf of Mannar- The coastlineis sandy with boulders and platforms of compressed sand stoneswith rough and uneven surfaces ocouring at different placeswith luxuriant dense growth of algae (Plate 1 A, B,C;&‘vD.).The present sporulation study was made by collecting theplants fortnightly during the:spring tide periods from threestations in the Gulf of Mannar side namely Thonithurai,Pudumadam and Kilakkarai (Fig.1). From Kilakarai (79O47'E,
9°121N) tetrasporic plants of Gelidiella_ggggg§g (Forsskal)Feldmann|g: Hamel were collected from the reef situated in thesubtidal region and oystooarpic plants of H1Dnea_musCiformiS
(Wulfen) Lamouroux from the rocks in the intertidal region.Collections of tetrasporic and oystocarpic plants of Graoilariacorticata J. Agardh and Hypnea musciformis were made from therocks in the intertidal region at Pudumadam.
Tetrasporio and oystocarpio plants of Graoilaria ggg;;§(Gmelin) Silva were collected from the subtidal region atThonithurai (79°E, 9O16'N). Plates (2-8) show the photographsof the specimens of the above four red algal speohes. Thematerial collected from all these stations were brought to thelaboratory in polythene bag containing sea water and theywere used for spore liberation studies.
Figure No.1 Coastline of Mandapam showing the
three collection localities namelyThonithurai, Pudumadam and Kilakkarai.Inset is the map of India showingMandapam Region.
mmcmm;. gnu mxo§zm «xmmm norrmnfioz ronptjmm
_
.0 b P .4 M Nmvh ... w.
lmo.
_vCOCIb0_....I
£DZODflD..£
_1IOZ..fiICm....._
xwrpxxpmb.
To.
uh i A 2 .3 h .._M
F mo+ 1 mo.
_“o. ,3. uo.
Plate No.I (A~D) Thonithurai coast showing bouldersand platforms of compressed sand
stones with seaweed growth.
,r.«v-giver»-var-»~o—-4re~. >.‘- .4
PLATE-I.
30
3.1 SPORE QUTPUT
Tetrasporic thalli of all the four red algae and cystocarpicthalli of Gracilarig'gg§£;ggg§ andl§.;ggg;;§'were used inexperiments carried out to study the seasonal changes inspore output, diurnal periodicity and also the effects ofenvironmental factors. Cystocarpic plants of Hypneamusciformis were not available in all months of the year.Depending upon the availability of carposporophytes,information was collected on seasonal and diurnal aspectsof carpospore shedding of this species. However, experimentson the effects of selected environmental.factors on carposporeoutput of Hypnea musciformis were conducted during themonths when cystocarpic plants were available.
Small clumps of Gelidiella acerosa with well developedstichidia and 5-6 cm long fronds of Gracilaria corticata,‘Q. edulis and Hypnea musciformis with well developedtetrasporangial sori and mature cystocarps were used foreach spore shedding experiment. The materials thusselected for the experiments were cleaned and washed severaltimes in sterile sea water. The thalli bearing fertileramuli or stichidia of Gelidiella acerosa were placed inpetri dishes of 5 cm diameter containing 20 ml of sterilesea water (Plate 9A) and fertile (tetrasporic and cystocarpic)
31
fronds of Gracilaria corticata,_g. edulis and_fl1pggmusciformis in 9 cm diameter petri dishes filled with 50 mlof sterile sea water (Plate 9B & 9C). For estimating the
spore producing capacity in the laboratory at roomtemperature the experimental sets were illuminated with alight source of 500 lux for 8 : 73 LD cycle(Plate 9A,B & C)For collecting the information on the seasonal spore outputthe spores liberated in the petri dishes were counted after24 hrs every day and for diurnal spore output, sporesliberated into the petri dishes at different times of theday i.e. at 4 hrs intervals from 2 PM onwards were counted.Tetraspores and carpospores liberated in the petri disheswere counted, as given below :
3.2 SPORE COUNT
At the end of each experiment, the liberated spores in thesea water in each petri dish were mixed thoroughly with afine brush and the spore suspension transferred to ameasuring cylinder (50 ml, 100 ml or 250 ml). The volume
of the spore suspension was adjusted to 20 or 25 ml forGelidiella gperosa and 50 or 60 ml for Gracilaria corticata,
‘Q. edulis and Hvpnea musciformis using washings of the petridish and sterile sea water. When spore shedding was high,the spore suspension.was diluted upto 50 ml or 100 ml or250 ml. A sub sample of 1 ml of the spore suspension was
32
then pipetted out into a plankton counting chamber andthe spores present in all the squares of the chamber werecounted with the help of hand tally counter under amonocular microscope. The degenerating sporeswere notcounted. From the mean value of two counts and the total
volume of the spore suspension, the spore output from thefruiting material was estimated. Depending on theavailability of the tetrasporic and cystocarpic plantstwo or four experiments were conducted separately in amonth for collecting data on seasonal and diurnal sporeoutput. Fresh weight of the fronds of these four red algaewere noted at the end of each experiment for computingtheir spore output per gram fresh weight of the plant. ForGglidiellg acerosa fresh weight of 5 clumps and the totalnumber of stichidia occurring on each clump were recorded"every time to estimate the number of reproductive ramuliiper gram fresh weight of the plant material. Theprocedures followed for seasonal and other studies onspore output are outlined as given below :
33
3.3. SEASONAL_§HANGES IN SPORE OUTPQE :
Every month four experiments were conducted with Gelid ellaacerosa, Gracilarga cortioata,5§. edulis and Hgpgea musciformisfrom October 1981 to September 1983 to understand the
seasonal changes in sporulation. The tetraspores andcarpospores liberated daily (every 2h hours) till the plantsdischarged spores upto a maximum of 30 days were counted to
lcnow the variations in the output of tetraspores andcarpospores in different days under laboratory conditions.The total number of spores liberated at different days fromthe plants collected every month was used for estimatingthe seasonal changes in the spore output and it is expressedas spores per gram fresh weight of the sporulating thalli.
3.1+ p_:g_U_13N_;A_g PERJODICITY IN §1PoRE. .oL1rsPAU$ :
Information on the diurnal changes on spore shedding wascollected by changing the material at every four hoursfrom one petri dish to another containing sterile sea water.These diurnal changes in spore production were studied for
two years from October 1981 to September 1983 for thetetraspores of Gelidiella acerosa, and for the tetrasporesand carpospores of_Grac1laria oortigata and.§. edulis andHxpnea musciformis. The diurnal rhythm in the liberationof oarpospores in Hypnea mggciformis was observed for oneyear only from August 1982 to July 1983 from Pudumadam area.
34
Data could not be collected on the diurnal periodicity oftetraspore output in Gelidiella acerosa, Gracilaria corticataand Hgpnea musciformis for the month of December, 1981.
The spores liberated at four hour intervals i.e; between 2 PMand 6 PM 3 6 PM and 10 PM 3 10 PM and 2 AM ; 2 AM and 6 AM-;6 AM and 10.NM and 10 AM and 2 PM were counted as stated
earlier and the data are presented as the percentage of sporesper gram fresh weight of the fruiting thallio The materialswere collected during the spring tide period of every monthand both seasonal and diurnal experiments were commenced
at 2 P; These experiments were conducted at room temperaturein the laboratory (29 3 3 °c) and at light intensity of500 lux during the day time for 8 hours from 10 AM to 6 PM.i.e. 8 i T6 LD photo regime.
3.5 EFFECTS OF ENVIRONMEflTAL FACTORS ON SPORE OUTPUT :
For assessing the effects of various environmental conditions3-9 experiments were conducted for each factor. The methodsadopted for each factor is described in the followingparagraphs;
35
3.5.1 EXPOSURE TO AIR on DE_$ICCATION :
For studying the influence of desiccation or exposure toair, fertile fronds of §gl;g;g;;g_agero§a, gracilarig_gorticata,_Q. edulis and-Hxpnea musciformig were exposed toair in the laboratory. Before exposing to air, the materialswere washed off any foreign particle and then blotted withthe blotting paper. The materials exposed to air uptoeight hours in the laboratory (from 15 minutes, 30 minutesand 1 hour intervals) were then transferred to the petridishes containing sterile sea water. Fronds that were keptthroughout the period of the experiment in submerged conditionswere used as controls, The spores liberated were countednext day after 24 hrs. While conducting these_experiments,the laboratory room temperature varied from 26.5 °_to 34.3 °Cand the relative humidity varied from 50 % to 87 %. Therelative humidity was found out with the help of a hygrometergThe experiment was conducted under a light intensity of500 lux and 8 :'T5 photo regime.
36
3 .5 .2 SALINITY :
A stock solution of 100 o/oo salinity was prepared byevaporating the sea water collected from the inshore area.of Mandapam and estimating the salinity following theWinkler's volumetric method of Strickland and Parsons(1968).The stock solution was sterilised and used for experimentalwork. From this stock solution the lower salinity gradeswere prepared by adding requisite quantity of distilledwater. The spore output in the four algae was estimatedat 0, 1o, 20, 3o, 40, 50, 6o, 70, so, 90 and 100 o/oosalinity by maintaining the petri dishes at room temperatureand providing 500 lux day light fluorescent illuminationfor 8 hours and 16 hours dark cycle;
3 .5 .3 L;LGH;TL IN§f'EN§I§:Y 3
Light intensity experiments were carried out at 6 differentilluminance values (0, 500;‘1000, 2000, 3000 and 4000 lux)and the petri dishes containing fruiting materials weresubjected to 24 hour light cycle for sporulationy
3.5 .4 _E1-I_C_)_TOPERIOD :
The combined effect of three different light intensities
(500, 2000 and 4000 iux) and day length on spore sheddingof the four red algae were studied. The petri dishescontaining the fruiting fronds (tetraspores and carpospores)
immersed in sterile sea water were subjected to 24 hoursdark (0 : EZ) and light (24 :'5) cycle with 4 hours intervali.e. O : EZ, 4 : E5, 8 : Z5, 12 : TE, 16 : 5, 20 : Z and24 : 5 using three light intensities.
3 - 5 -5 .lE;“[email protected],.E-..i/§.§E.‘..lR.-‘$3.. 3
JThe influence of 10 different temperatures-15 °, 0 CC, 5 °C, 20 °c, 25 °c, 30 °C, 35 °c, 40 °c, 45 °acand 50 °c on spore shedding was studied by maintaining thepetri dishes for 24 hours in a temperature controlled darkeven or refrigerator or deep freezer, In all these experiments on different factors the spores liberated in the petridishes were counted after 24 hours and the spore output wasexpressed as spores per gram fresh weight per day,
3.6 GERMINATIoN;oEgseoRa§ 2
While counting the spores in various experiments plannedin this study, data on the germinating spores were alsocollected separately. The frequency of these germinatingspores (within 2# hours of their liberation) in different
38
-months of the year and also in the experiments conductedto study the effects of exposure to air, salinity, lightintensity, photoperiod and temperature on spore shedding.
3.7 SEASONAL QHANGES IN TEE HYDROGRAPHICAL PARAMETEg§ :
In an attempt to determine the seasonal fluctuations inthe hydrographical parameters in the localities from wherematerials have been obtained, fortnightly data were
collected from October 1981 to June 1983 on the surface seawater temperature, pH, salinity and dissolved oxygen, whilecollecting the materials. The surface sea water temperaturewas measured and the atmospheric temperature in the fieldwas also recorded simultaneously. Water samples werecollected from each collection spot for analysis of pH,salinity and dissolved oxygen. For estimation of dissolved
oxygen, the water samples were fixed with 1 ml of MnCl2(Winkler A) and 1 ml NaOH + KI (Winkler B) solutions
immediately after collection on the field. In thelaboratory, pH of the sea water collected from the workspots was found out with a pH meter. The salinityand dissolved oxygen of the sea water were estimated bystandard methods given by Strickland and Parsons (1968).
39
.CH§§TER - 4
RESULTS
Results obtained on the fruiting behaviour, dailyand monthly changes in spore output, diurnal periodicityin the production of spores and the effects of environ*mental factors such as desiccation, salinity, light(intensity and photoperiod) and temperature on spore
shedding in the four red algae of‘Mandapam coast namely§g;;;e;lg_ggg£g§g, Gracilaria corticaga,.§..gggl;§ and§ypggg'gugg;£g£mi§ are presented here. The datacollected on hydrographical aspects such as temperature,
salinity, pH and dissolved oxygen and also atmospherictemperature recorded from the three collection localitiesnamely Thcnithurai, Pudumadam and Kilakarai are alsopresented in this chapter.
4.1.1 _]_§‘_RUI'._[flI\IG BE1I_j_._€_1_’\I_lOUR ’:
Population of Gelidiellg gcerosa_at Kilakarai, ggggglgggggggigagg at Pudumadam,l§. edulis at Thonithurai andtHypnea musciformis at Pudumadam and Kilakarai in the
vicinity of Mandapam occurred throughout the year.Igfiraspgric plants of all these four species were foundcontinuously from October 1981 to September 1983.
CvS’°°°m°1“I5—*=-0 Plants of .9_<2r.ti.9.a39s and .<‘:~ .e~:1.9_;ie
40
occurred in all months of the year. During the two yearperiod October 1981 - September 1983, carposporophytes
of‘§ypgg§;my§ciformis at Pudumadam occurred only for13 months from August 1982 to July 1983 and September
1983 and at Kilakarai except in November and December
1981 and January and June 1982 they occurred in theother twenty months. Cystocarpic plants of gglidiellaacerosa were not found during the entire period of thestudy.
4.1.2 gQflEflQLOGICAL APPEA§§NCE OF THE FRUITING
EfiOND$ Qf THE FOUR"SE§g;§§mpF AGARQE§gTE§ :
The identification of cystocarpic and tetrasporicthalli in the field with the ordinary eye is mostdesirable for one to be able to facilitate the processof collecting the fertile thalli to be used in thesporulation studies.
The "fruiting" thalli of the four algal species,are shown in plate (2 - 8) and figures, (2E - 2H).The tetraspores are produced by a tetrasporic thallusand carpospores by a cystocarpic frond.
Cystgggrpic Plants
In all the species described the carpospores areproduced in swollen sessile hemispherical structures
41
with dark red, brown contents, known as "cystocarps"which are borne singly or in an aggregation of two,three or five cystocarps. These occur all over thesurface of the thallus and also close to the thallusapex. The cystocarps observed in Gracilarig cortigatg,G. edulis and gxpggg musciforgig vary in size and mayattain a diameter of up to 0.5 mmnto 0.9 mm and 4.0 mmto 1;2 millimeters. Cystocarps are (Fritsch, 1945)part of the carposporophytic generation which develops
yparasitically on the female gametophyte. The morphological appearance of the cystocarpic plants is shown inplates 2 - 7 and Figures 2F and 2G.
Tetrasporic Plgggs
The tetrasporophytic thallus producing the tetrasporesin Gelidiella ggggggg, Gracilaria gg££$cggg,‘§.‘ggEl;§andiflypggg muscigprmis are shown in plate (2-7) and
(Figures, 2E & H). In Qelidiel;g‘§gggg§g and‘§ypgg§_gusciformis a fertile tetrasporophytic plant isdistinguished on the basis of its swollen side branchlets which are then called as stichidial branches(Tanaka 1941) and from the presence of lack of anyhemispherical swellings. The swollen region of thestichidia or ramullii, where the tetrasporangia areconcentrated, is usually of a darker colour (Pink
42
brown in case of Gelidiella acerosa and darker brown incolour in.Hypnea ggggigprmis) than any portion of vegetative branchlets (Pate 2 & 7) and Figures 2E and 2H. In
Gracilaria corticagg the tetrasporangia are found allover the thallus just below the cortex and embedded inthe cortex layer. The thallus is flatterned as shownin.Plate (3A and AAB) with numerous dichotomous branches
arising from the basal attachment. In_Q. edulis thethallus is cylindrical in structure, dark red, brown,green in colour with numerous fronds arising from thebasal disc (Plate 5A.& 6A). The tetrasporangia are
borne on the tetrasporic fronds just below the cortexlayer. In these species there is no specialised appendage as is found in Gelidiella acerosa and Hypneamusciformis.
Thelgglggigllg acerqgg plants are thick wiry, mostlylight brown dark brown to green in colour. The frxmndsare stiff, cartilaginuous show repeated pinnatebranching (Fritsch, 1945) (Plate 2A, B and Figures 2E)
Hypnea Musgggrmis
(Plate 7A, B and Figure 2H) has fleshy terete thalliwith numerous long and short branches and are incurled atthe tip, with dark green, brown in colour.
Plate No.11 A Qg}idie1la acerosa, habit.(B-C) Tetrasporic thalli Q, acerosa
t»:i"bh (s- stachiciia,B-branch, bl- L-manchletl 'T— H101“marl-zed an-ow) .
PLATE _II.
Figure No.2.E Fruiting frond of Gelidiellaacerosa, showing tetrasporangialstichidia. Arrow showing T-thalli,S-stichidia, b-branch, bl-branchlet.
F|GURE..2E1ond 2‘hI
ID-.J
‘II
Plate No 111 A Gracilaria coxticata (t9tTaSP°ri°)habita
B Ea ..C.<2.r;1;,i.._<::«:;:;; CCvS*~’~°°a“Pi°9 }“"‘b‘“"
PLATE- .
2.
%§€§§&_'§é§¥ ,&fiflfififi
Plate No.IV A kGr-acilaria oortioata (tetrasporic)habit.
B Tetrasporic thalli of Q. 'c.or'l:icatafmagrfified) .'_i.-.:_'_: A‘
C Cystocar-pic thalli of _g_. corticata.D Cystooarpic thalli of Q. corticata
showing oystooarps. (C _ma,,,<ed C)arrow)
PLATE_lV.
9..
104»Jun.
.C
VVs!
:I
7!....
Figure No.2.F Cystocarpic frond of Qgcggiarig_c_:_gg§;gg3g showing the arrangement
of cystocarpa. Arrow showingC-cystocarp, T-thalli, b-branch,bl-branchlet, X 10.
FIGURE- 2 F
Plate No9V A Tetrasporic plant of Graoilaria. Ig§g;;§3 habifieB Cystocarpio plan? of‘§. edulis, habit.
PLATE-V.
Plate No.VE AB
C
Tetrasporic plant of firaci ‘a edulis.Cystocarpic thalli of‘Q. edulis.Cystocarpic thalli of'§. edulisshowing cystocarps.
PLATE_V|.
Figure NO.2.G cystocarpic frond of Q.showing the arrangement of
cyatocars. Arrow showingC-cystocarp, T-thalli, b-branch,
-26, qondz
C‘~>
Plate No .VIf[ Hxpneg muscizformis, habit.
PLATE- VII
Plate No.VIII A Tetrasporic plant of Hzpnea musciformisB Cystocarpic plant oflgyphea
musciformis with oystocarps.
PLATE _V|||.
Figure NO 02 oH Part or a tetrasporic frond of g1pgg__a_
%gg9_;_'x_z,1__i__s to show morphological
appearance of tetrasporangial stichidia.Arrow showing T-thalli, S-stichidia,b-branch, bl-branchlet.
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PLATE -9
.. l.,_ .
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Plate No.X A Tetraspores of Gelidiella aoerosa.B Spores of Gracilgggg corticata.C Tfiaores of ~_G_;r‘__Qéi;a£_,:_I:a ‘egigig.
D Spores of.§1Qg§g_musciform;s.
PLATE- X.
43
4-2 §§.9.8.E;oUTPiUE =
Tetraspore output at different days during variousmonths of the years were studied in Gelidiella acerosa,Graoilaria oorticata,Ԥ. edulis and Hxpnea musciformisare presented in Tables 1-4 and that of carpospores in
Graoilaria_gorticata,.Q, edulis and‘ggpggg musoiformisare given in Tables (5-8) to show the trend in thedaily liberation of spores. In the four algae studiedmaximum shedding of both tetraspores and Carpospores
were seen on the first day almost in all the months.The tetraspore output decreased from 2nd day onwards(Tables 1-4) whereas carpospores were liberatedrhythmically with peaks at intervals of differentdays in many months.(Tables 5-8). To show the rhythmicliberation of carpospores in gracilaria corticata,w§.edulis and‘§1pgg§ musoiformgg, the data for few monthsfrom Tables 5 to 8 where this trend is clear are plottedin Fig. fi&3.Uhder laboratory conditions tetraspore output was seen from 6-14 days in.Qg;;g;gll§_ggggg§§,6-27 days in graoilaria oorticata, 3-30 days in Q, edulisand 3~23 days in Hgpnea musciformis during differentmonths of the year (Tables 1-4), Shedding of carposporeswas found for 6—3O days in.Q3gg;;ariapgorticata, 10~3O
days in G. ggulgg and 2-24 days in.§zpQgg muscifoggig0&1
Figure No.2 Seasonal variation in stichidia (A)and tetraspores (B) o£'§§;gg;g;;§ageroaa and tetraspores (C) andarpospores (D) of Grac;1ar;a coggicat .
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during the period of this investigation.CTables 5-8).‘ Plate(10) shows the photographs of tetraspores and carpospores ofthese four red algae.
4.3 ,SEASONAL CHANGES Ifim§PQfi§_QUTRflT
The total quantity of tetraspores and oarpospores liberatedat different days from plants collected during every monthewes used for estimating the monthly variation in sporeproduction. "The number of stichidia and tetraspores pergram fresh weight of the'plant of Gelidiella_acerosa fromOctober 1981 to September 1983 are given in-Fig. 2. Thetetraspore and carpospore output for Qracilarie oetticataare plotted in Fig. 2c,%.ncz for Gracilaria e_dul_i_§ and g1_p_g_e_g_
musciiormis in Fig. 4.
In_Qglidiellg‘ggg£g§§ stichifiia containing tetrasporeswere found in all months of the year without any seasonalvariations in their abundance. However, maximum number ofstichidia per gram fresh weight of the plant occurred
between.Apri1 and July 1983. During the period of thisstudy, the number of stichidia varied from 13 to 418 pergram fresh weight of the plant (Fig. 2A). Figure 2B showsthe monthly and yearly variations in tetraspore sheldingor qg}idie1l§.2£££gg§. Spore liberation occurred in allthe months of the year. Though there was no regular trendin spore output, maximum shedding of tetraspores was observed
Figure No.3 Rhythmic liberation of carposporesin Gracilaria corticata (A), Q.edulis (B) and Hxgnea musciformisfrom Pudumadam (C) and Kilakkarai (D).
CARPOSPORES / G. FR. WT. (XIOOO)
FIGURE - 3 L50.; GRACILARIA CORTICATAA ‘ . *—° Januar I983-1 ‘ 0-0 April I 83I51 \‘ - ' e \\ . A ‘ A''—¢ JUU I9'. . ‘ \IO-+ O ' J X X54 , . . \\ ‘ _ A .T A ° ' ' ~ ‘ . . .0‘ _ __, .1-_-_.50* - GRACILARIA Eouus'* w-0 December I98230'‘ r——-0 April I9831; . , . ‘ ‘ A'—‘—A A0908? I983‘ I . A °3. .-4 . .~ V -1of y u .- ‘ HYPNEA uuscnronms90* . X (Pudumadam)‘ <>——o Navembe I982504 °\ - "“‘ January I983W ‘x, _._ . A"“¢June I983IO; ‘V. . . I-4 .4. ‘-4 C - ' ‘ I:o , 4--- — -_ ..a e -ea
90" HYPNEA MUSCIFORMIS.. , mlauarai)50% ' or-0 Januacy I983I24’ , _ o—« June use:- - A——A August I9838-44 - ‘
.. D0 i f V 1 Y Y ' Y * I 'O
45
in the first year in the month of July and in the secondyear from April to September with a low value in July isshown in Fig. 2B. The tetraspore output ranged from 5,744to 3.89.793 spores per gram fresh weight and there was norelationship between the number of stichidia and sporesliberated from the plants collected in different monthsof the year in Gelidiellg acerosa.
Shedding of tetraspores and carpospores in Gracilariacorticata were found in all months of the year and regularchanges were not observed either in the output of tetrasporesor carpospores (Fig. 2C and 2D). The tetraspore outputvaried from 8,181 to 92,277 and carpospores from 8,086 to
h,87,178 spores per gram fresh weight. In_§racilaria3ggg;i§also tetraspore and carpospore output occurred in all themonths of the year without any marked seasonal fluctuationsin the production of tetraspores and carpospores (Figs. 4Aand 4B). The tetraspore output varied from 2,135 to 1,09,565and carpospores from 28,652 to 3,27,833 numbers.
Data on tetraspore output was collected for all the monthsof the year for Hypnea musciformis occurring in theintertidal area at Pudumadam (Fig. 4C). Information oncarpospore output was gathered only for 13 months fromAugust 1982 to July 1983 and September 1983 for Hypneamusciformis also collected from Pudumadam and for 20 months
Figure No.4 Seasonal variation in the liberationof tetraspores (A) and carpospores (B)of Gracilaria edulis and tetraspores(C)and carpospores (D) of Hzgnea musciformis
CARPOSPORES/q.h'.w!. TETRASPORES./g.fr.M. CARPOSPORES/q.?.'.w*. TETF<:-.SF'OF?ES/9 1’? W’.
lwbuH£.4 GHACILARIA EQJLISI<)(>()L)Lv
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for the same species collected from Kilakarai (Fig. 4d),since in the other menths oystocarpio plants were notavailable in the field. Marked seasonal changes were notseen in the liberation of both tetrspores and carpospores.The tetraspore output varied from 2,662 to 5,94,953 spores“per gram fresh.weight per day. The earpospore outputranged from 16,t12 to 6,46,585 and from 2,814 to 2,83,745
spores per gram ireshrmeight per day in.H1EneaJgg§g;§g§g;§_collected frem Pudumafiam and.Kilakarai respectively.
4.4 DIURNAL paelcnmcieew N the secne SHEDDING
To show the general trehdfi in the daily periodieity in theliberation of tetraspores and carpospores in differentmonths of the year, the mean values (expressed as percentage)of the experiments congested in each month for 2 years from_October 1981 to September 1983 with tetrasporophytes of
Gelidiella acerosa, Gracilari§%cortipata,4§.‘3§E;i§ andHzgnealmusciformis are plotted in Figs. 5 to 12. Datacollected on seasonal changes in the diurnal periddicityin the shedding of carpospores for 2 years from October1981 to September 1983 with Gracilaria corticata and_§.edulis. For 1 year period from August 1982 to July 1983with.HxEnea musciformis collected from Pudumadam (Station II)
and for 20 months 10: carpospore output in_HgEnea musciformiscollected from Kilakkarai (Station III) are plotted inFigs. 13-17.
Figure No.5 Diufnal periodicity in the sheddingof tetraspores of Gelidiella acerosacollected from October 1981 to
September 1982.
°/o TE TRASPORES/ g. fr. wt.
FIGURE - 5 - GELIDIELLA ACEROS/-\
9o ocToBER’8I APRIL
454/)\' MQ 1 % n '90" NOVEMBER MAY
45— J:_90- rDECEMBER JUNE45‘ NO DATA
90‘ JANUARY '82 JULY
45q —A—‘-J90‘ FEBRUARY AUGUST
45- / \, M90‘ MARCH SEPTEMBER
2PMI
6 fo zkuésTIM E
IO 2PM 2PM 6 ‘IO 2AM 6 IO 2PM
Figure No.6 Diurnal periodicity in the sheddingof tetraspores of Gelidiella acerosacollected from October 1982 to
September 1983.
°/o TE TRASPORES/g.fr.wt.
F|GURE-6. GELIDIELLA ACEROSA
90“
45
90
45*
90‘
45‘
1
90
45fi
90*
45*
90
45"
OCTOBER'82 APRK.
NOVEMBER MAY
DECEMBER JUNE
JANUARY '83
FEBRUARY
AwJULY
AUGUST
MARCH SEPTEMBER
T I2AM 6IB HDZPM _ w 1 T
ZRM 6 HDZAM 6 IOZWMTIME
Figure No.7 Seasonal variation in the diurnalperiodicity in the liberation oftetraspores of Gracilaria corticatacollected from October 1981 toSeptember 1982.
FIGURE--'?‘. GRACILARIA CORTICATA60-+ OCTOBER'BI APRIL
30”-Hr_g..._.....fl=: "E-. » -~ ______,,__..\. ' #_______. ‘ " I ' '50”“ NOVEMBER MAY
-3'3 6°" DECEMBER JUNEC»N. 30*-w NO DATAU)M0: .O 60-- JANUARY 82 JULYca.0) <10:l-uJ 60“,‘_ FEBRUARY AUGUST
+~-+;fr-===9'“""fl 3 3‘ *‘6°” MARCH SEPTEMBERso-—«
°‘“‘“‘i"“‘iF.:’rM I I I I 1 | I2PM 6 no 2AM 6 no 2PM 2PM 6 I0 2AM 6 IO 2PM
TIME
Figure No.8 Seasonal variation in the diurnalperiodicity in the liberation oftetraspores of Graoilaria oortioatacollected from October 1982 to
September 1983.
FIGURE- 8. GRACILARIA CORTICATAso-Y OCTOBER'82 APRIL30- \"\.,
.,..a/501 NOVEMBER MAY
30-‘ -—q.._- 60‘ DECEMBER JUNE3"5. 3°?D\
cnLL] _(1 6° JANUARY '83 JULY00. 30-4<1:
<12:1: — .5 5°‘ FEBRUARY AUGUSTF
60-+ MARCH SEPTEMBER
1 I I V2PM 6 I0 2AM 6 IO 2PM 2PM6lO 2AM IO 2PM
TIME
Figure No.9 Seasonal changes in the diurnalperiodicity in the liberation oftetraspores of Gracilaria eduliscollected from October 1981 to
September 1982.
°/0 TETRASPORES/g.fr.wt.
F|GURE.9 GRACILARIA EDULIS90- OCTOBER'8l APRH.45-‘ >9°“NovEMBER MAY45‘
9°“ DECEMBER JUNE45’ W90- JANUARY'82 JULY
45~ 9-—4—/—X¢90- FEBRUARY AUGUST
45— o—+~9//—\9°“ MARCH SEPTEMBER45-*0-‘ I I I I
2PM 6 IO 2AM 6lO 2PMI I I I I
ZPM6 IO 2AM 6 IOZPMTIME
Figure No.10 Seasonal changes in the diurnalperiodicity in the liberation oftetraspores of Ggggilarggjggggygcollected from October 1982 to
September 1983.
°/0 TETRASPORES / g.fr.w1.
F|GURE.10. GRACILARIA EDULIS
9°‘ OCTOBER'82 APRH_
45_ O=i?9°‘ NOVEMBER MAY. ,.9°‘oEcEMaER JUNE45d90-J , “JANUARY 83 JULX
45‘ ._\.—_././‘\‘/—¢90 FEBRUARY AUGUST
45- :O—¢———A90"‘ MARCH SEPTEMBER45‘
0 . ‘ I .r*#b‘f’7/fir/:\j2PM 6 IO ZAM6 I0 2PM ZPM6 IO ZAM6 IO 2PM
TIME
Figure No.11 Seasonal variation in the diurnalperiodicity in the shedding oftetraspores of Hzgnea musciformiscollected from October 1981 to
September 1982.
FIGURE -11. HYPNEA MUSCIFORMIS
90"‘ OCTOBERx8| APRIL
45R90-4 NOVEMBER MAY
*_: 45-13 32>/Z\\,:_" .¢9o~\ DECEMBE JUNEU) 45
Ld >——¢—4/Z\\oO:O _0- 90- 0') JANUARY 32 -JULY<1
E 45LIJI-'
90"‘9 FEBRUARY AUGUSTo\
45d 47%90" MARCH SEPTEMBE45-‘
0 I I I I I I I I2PM 6 IO 2AM 6 I0 2PM 2PM 6 I0 2AM
I I6 IO 2PM
TIME
Figure No.12 Seasonal variation in the diurnalperiodicity in the shedding oftetraspores of Hypnea musciformiscéllected from October 1982 to
September 1985.
F|GURE-12. HYPNE/-\ MUSCIFORMIS90- OCTOBER'82 APRIL
45
_9°‘ NOVEMBER MAY
:45T o——--:——//\0'»
\9°" DECEMBER JUNE(1)LLJ45-1
o0.
2 90- JANUARY'83 JULY01I-45LLJF fi 1 L
90‘ FEBRUARY AUGUSTo\"45
90- MARCH SEPTEMBER45‘0‘ I I I T I I I
ZPM 6 IO 2AM6 IO 2PM2PM6 IO ZAM6 IO 2PMTIME
‘47
In ggligigllg acerosa except in December 1982, in allother months peak output of tetraspores was observedwithin the first hours after commencing the experimentsi.e. between 2 PM and 6 PM. After 6 EM there was a sudden~
fall in the liberation of tetraspores and the percentage ofspores liberated was minimum from 10 PM to 2 PM (Figs. 5 & 5).Sudden shedding of tetraspores with prominent peak at oneperiod of the day was not seen in different months of the
year in Gracilaria gprticata,]§._edulis and Hypneagmusoiformis(Fig. 7 to 12). In general, the quantity of sporesliberated was more during the period of the day from 10 PMto 2 PM and minimum quantity of spore were liberated between2 PM and 10 PM in all these three species of the orderGigartinales. Similar observations with maximum carposporeoutput from 10 PM to 2 PM were made from the cystocarpic
plants of Gracilaria cortigata and4§. edulis (Figs. 13 to 16).But in.Hypnea musciformis maximum liberation of oarposporesoccurred either from 10 PM to 2 AM or from 2 AM to 6 AM
during the one year period from August 1982 to July 1983collected from Pudumadam. The same was also observed for
2 years in the carpospore output of Hypnea musciformiscollected from Kilakarai. The quantity of oarposporesliberated between 2 PM and 10 PM in Gracilaria cortigata,Q. edulis and Hypnea musciformia was less (Figs. 13 to 17)as observed in the tetraspore output of these three species.
Figure No.13 Diurnal periodicity in the carposporesoutput of Gracilaria corticata collectedfrom October 1981 to September 1982.
g.fr.w1. 8 3
°/o CARPOSPORES /
FIGURE. 13. GRACILARIA CORTI CATA
60
aoa
0‘ 0
30-1
OCTOBER'8| APRK
NOVEMER
W/’\DECEMBER JUNE
60"
30
60‘
30*
60
30*‘
JANUARY'82 JULY
FEBRUARY AUGU ST
{L
MARCH
0.: I I T
2PM 6 IO 2AM 6 I0 ZPMZPM 6TIME
SEPTEMBER
,6 +‘fiF‘#//K/i\\r\‘IO 2AM 6 IO 2PM
Figure No.14 Diurnal periodicity in carposporeoutput of Gracilariaicorticatacollected from October 1981 to
September 1982.
°/o CARPOSPORES / g.fr.wt.
01 0g R
FIGURE .14. GRACILARIA CORTICATA
6°‘ OCTOBER '82 APRIL
30
50* NOVEMBER MAY30*
6°" DECEMBER JUNE30-‘
6°" JANUARY'83 JULY30“
6°‘ FEBRUARY AUGUST
5°‘ MARCH SEPTEMBER
W I T 7 I I I I I I I2PM 6 I0 ZAMG IO 2PM2PM 6 IO ZAM6 IO 2PM
TIME
Figure No.15 Diurnal periodicity in the sheddingof carpospore of Gracilaria eduliscollected from October 1981 toSeptember 1982.
°/o CARPOSPORES / g.fr.wt.
F|GURE-15. GRACILARIAJEDULIS
90- OCTOBER '8! ApR||_
45‘ ¢—4—9°‘ NOVEMBER MAY
45- 0-=f/X9°‘ DECEMBER JUNE45‘90-‘ JANUARY'82 JULY
45"
9°‘ FEBRUARY AUG'UST459°‘ MARCH SEPTEMBER45o
2PM 6 IO 2AM 6 IO ZPMZPM 6I I T T I
I0 ZAM6 IO 2.PMTIME
Figure No.16 Diurnal periodicity in the shedding ofcerpospores of Gracilaria eduliscollected from October 1982 to
September 1985.
90+ OCTOBER'82 APRIL
°/.. CARPOSPORES / g.fr.wt.
F|GURE-16. GRACILARIA EDULIS
45-‘
_a-i-—/./.§‘\.90- NOVEMBER MAY4? u:I
9°‘ DECEMBER JUNE45- f\\,90- ,JANUARY 83 JULY
45-w &-::’Q—/\90? FEBRUARY AUGUST
45- r—%%—/\904 MARCH SEPTEMBER45-‘
O I V ' ' 7 I T I T2PM 6 IO 2AM 6:10 2PM2J'PM 6 no 2AM 6 I0 2PM
T I hfl E
Figure No.17 Diurnal periodicity in the liberationof carpospores of.Hypnea musciformiscollected from August 1982 toJuly 1983.
F|GURE..17 HYPNEA MUSCIFOMIS60; AUGUST '82 FEBRUARY30- f\60- SEPTEMBER MARCH
81
O? O 1 OCTOBER APRIL
5°“ NOVEMBER MAY
CARPOSPORES/gfiwtO’) 04 0| (1) A ‘I’ ? DECEMBER JUNE
o\ 30-«
60"JANuARv '83 JULY30
0 1 I I I I 1 I I T I2PM 6 no 2AM 6 |O’~2PM 2PM BIO 2AM 6 no 2PM
TIME
48
#05 §§§§§TS OF SELECTED ENVIRONMEN3§£ FACTORS ON
§£QBE SHEDDING
Results obtained on the effects of environmental factors
such as exposure to air and desiccation, salinity, light.and temperature on tetraspore liberation in Gelidiellaacerosa and tetraspore and carpospore output in Gracilariacorticata, Q, edulis and Hxpnea musciformis are given below:
4.5.1 §;<_PosURE TO AIR on DES;_CCATIO_l\_I
This experiment was conducted with a View to study theeffect of exposure during low tides and the resultantdesiccation of plants on spore production and also tounderstand the spore release from plants occurring inshaded areas in the field. Changes observed in the releaseof spores from Gelidiella acerosa, Gracilgria cortica;a,.§.edulig and Hxpnea musciformig in the controls (0 minuteexposure) and at different periods of exposure to air inthe room (Temp. 2835 to 30.7 °C and R.H. 52 % to 87 %)
are shown in Figs. 18 to 21.
In the experiments conducted with tetrasporic thalli ofGelidiella acerosa exposing them to air in shade in theroom at intervals of 15 minutes upto 120 minutes,sporulation was seen upto 105 minutes. .Maximum output oftetraspores was found in control i.e. in fronds submergedfor 24 hours and the number of spores liberated decreased
Figure No.18 Effect of desiccation on the tetrasporeoutput in Gelidiella acerosa.
F|GURE-|8. GELIDIELLA ACEROSA
XIOOO
I6-+
I4-4
E 1
51
0 1
TETRASPORES/gfnwt/day
CD I
O I I I T I I0 I5 30 45 60 75 90 I05 I20DURATION OF EXPOSURE (minutes)
Figure No.19 Effect of desiccation on the liberationof tetraspores and carpospores inGracilaria corticata.
FIGURE- I9. G RACILARIA CORTICATA
X2000
201% \;.\
l.4J
S 1
5 J
G I
SPORES / g.fr.w1./ day
41: m
1
0-—0 Tetrospores
A-—~A Corpospores
“ ‘T’ /1 I | I I I I Ié' -—I
0DURATION OF EXPOSURE
I
'/4 ‘/2 $4 1 1'/41'/2 13/4 2 2('/2 3 3')/2 4hours
Figure No.20 Effect of desiccation on theliberation of tetraspores andcarpospores in Gracilaria edulis.
F|GURE-20. GRACILARIA EDULISX2000
I4-*
F5 1
|0'+
O I
O5 1
SPORES / g.fr.w1. / day
A 1
‘--TetrcnsporesA‘ Acorpospores
5678I ‘T 1 II I '3 I I! I0 /211/222/2 33/24(hours)DURATION OF EXPOSURE
Figure No.21 Effect of desiccation on theshedding of tetraspores andcarpospores in Hzpnea musciformis.
FIGURE -2|. HYPNEA MUSCIFORM I SXIOOO
60
40 °——0 TetrosporesAr—-A Corpospores. JI
-W
‘W
N O
3 1
N 1
SPORES / g.fr.w1. / day
5 L
8.46- \/4_
2..
O1|'|“'|'l'iH| IO /4 /2 3 11/41/213/422’2 33/2 4DURATION OF EXPOSURE (hours)
49
with increase in the duration of drying of plants atroom temperature. The spore output declined rapidly in15 minutes exposure and thereafter gradual decrease in theliberation of spores was observed in exposure of upto105 minutes (Fig. 18). In the experiments conducted withtetrasporophytes and carposporophytes of Gracilaria
corticata exposing them to air at V4 and 92 hr intervalsupto h hours, tetraspore output was found upto 4 hours andcarpospore output upto 392 hours. As observed in theliberation of tetraspores in'§glidie1la acerosa, maximumquantity of tetraspores and carpospores were released in0 minutes exposure and the spore output decreased graduallyfrom Vh hr exposure onwards (Fig. 19).
In the asexual and female plants of Gracilaria edulisexposed to air at V2 and 1 hour intervals upto 8 hours,shedding of tetraspores was seen upto 8 hours andcarpospores upto 6 hours. The quantity of tetrasporesliberated from 15 minutes exposed thalli was found to bemore than control. From 1 hour exposure there is gradualdecrease in spore discharge though there is slight increasein the release of tetraspores from 3V2 hours exposedplants over 2, 2V2 and 3 hr exposed plants. But maximum
quantity of carpospore output in gracilagig edulis wasfound in control with gradual decrease with increase in
50d
the duration of exposure (Fig. 20) as observed in thetetraspore and carpospore release of Gracilaria corticata.In the tetrasporic and cystocarpic fronds of Hgpnea musciformisexposed to air at Vh and 92 hour intervals upto 4 hours,tetraspore liberation was seen upto 4 hours. The quantityof tetraspores released from the thalli exposed at differenttime intervals varied very much without showing any trendin the spore release.‘ But maximum liberation of carposporesoccurred in centrol with gradual decline except at hourexposure (Fig. 21).
4.5.2 SALINITY
Effects of salinity on spore output in.Ge1idie11a acerosa,_gracilaria‘go§ticata,‘Q. edulis and H1pnea.gg§§g£§§g§y§are shown in Figs. 22 to 25 respectively. Spore outputvaried markedly in different salinities of sea watertested from 0 o/oo (0, 10, 20, 30, 40, 50, 60, 70, so, 90and 100 o/oo). In all the four algae, release of sporesoccurred from O o/oo to 90 o/oo and there was no sperulationat 100 o/oo. Peak spore output of tetraspores was found atno o/oo in Geltidifiellpa _a_c_§_r_o_§__e_1_ and at 30 o/oo in Gracpilaria
_ggrticata,‘Q. edulis and Hxpneapgusciformig. Though peakspore output occurred at these two salinities, thequantities of spore liberated at 20 o/oo and 50 o/oo inthe four algae were also more than compared with therelease of spores in other salinities.
Figure No.22 Effect of salinity on the tetrasporeoutput in Gelidiella acerosa.
F|GURE-22. GELIDIELLA ACEROSA
X I000
I6‘
I1
I-\-J 1
TETRASPORE / g.fr.w1. / day
on 5 L J
0. .O I Y I 1 1 1 I ro 20 40 so 80 I00SALINITY (°/ac)
Figure No.23 Effect of salinity on the tetrasporecarpospore liberation in Gracilariacorticata.
FIGURE- 23. GRACILARIA CORTICATA
xuooofi201 1:I6-1 /1 0-°Te1rospores
A--43 Carpospores3 I
§ 1
5I
Q I
SPORES/ gfnwt/day
‘f’
O I 7 I I I V&o 20 40 so so I00SALINITY (O/co)
Figure No.24 Effect of salinity on tetraspore andcarpospore shedding in Gracilariaedulis.
XIOOO
SPORES/g. fr. wt. /day
FIGURE 24 GRACILARIA EDULIS
46
36-‘
N O‘)I
CD 03L,‘ J
\\
m 1
o———o TETRASPORES
£r—1AcARPosPoREs
SAL INITY ( °/co)
Figure No.25 Effect of salinity on the dischargeof tetraspore and carpospore inHggnea musciformis.
FIGURE 25 HYPNEA MUSCIFORMISXIOOO
5 0 ‘ o——o TETRASPORESA-—A CARPOSPORES
-I3 O L
()1 O I
N O l
SPORES/g. fr. wt./day
m S IALL
O I I T \ I 7O 20 40 60 80 I00SALINITY (°/co)
51
4.5.5 LIGHT INTENSITY
Figs. 26 to 29 show the quantity of spores liberated fromGelidiella acerosa, Gracilarig corticatg,.§. edulis andgxpggg musciformis in dark ( 0 light intensity) and at 5different light intensities ranging from 500 to 4000 lux(500, 1000, 2000, 3000 and 4000 lux). Spore output variedin different light intensities ranging from 0 to #000 luxand spore shedding occurred in all 6 light intensities.The spore output in the four red algae was less in dark(0 light intensity), 2000, 3000 and 4000 lux lightintensities. Peak discharge of tetraspores in.gg;;diel1aggggggg, and tetraspores and oarpospores of Gracilariacorticata and'Q. edulis was found at 500 lux. The quantityof spores liberated at 1000 lux was also high in thesethree species. In.H1pnea musciformis maximum liberationof tetraspores as well as carpospores was observed at 1000lux light intensity. More number of spores were releasedalso at 500 lux light intensity.
4~5o4 EEMEEQAEHEE
Changes observed in the spore output of Gelidiella acerosa,
10 different temperatures (-15 O, 0 O, 5 O, 20 O, 25 O, 30 O,55 O. 40 O, 45 0 and 50 00) are given in Figs. 30 to 33.
Gracilarig_ggrticata, G. edulis and Hxpnea muscif0rmi§ at
$IA~
In Gelidiellg gggggsg tetraspore liberation was not foundat -150 450 and 5000. Minimum quantity of spores wasireleased at O0 and 40°C and the spore output values obtainedat 5° and 35°C were also low. Maximum quantity of sporeswas liberated in the gelidiaceous alga between 200 and 35°Cwith peak at 25°C (Fig. 30).
In.gga9ilaria ggggigagg there was no spore release at -15°,0° and 50°C and spore output was minimum at 5° and 45°C.
Peak discharge of tetraspores was observed at 50°C andcarpospores at 25°C. The quantity of tetraspores andcarpospores liberated in other temperatures was low (Fig;31).
Shedding of spores in.g£g;;gg;§.gggl;§ was minimum at0°, 5°, 40° and 45°C and low spore output was found at 35°C.There was no spore output at -15° and 50°C. Though peak
output of tetraspore was found at 30°C and carposporeliberation at 25°C, the spore output values obtained fortetraspores at 20° and 25°C and for carpospores at 20°and 30°C was also equally high in graoilaria edu1is(Fig.32).
In Hxpneapmusoiformig discharge of tetraspores was notfound at -15°, 0° and 50°C and oarpospores at -150, 00,,#00 and 45°C; More number of tetraspores and carpospore
52
liberation was observed between 20 OC and 35 0C with peak
spore output at 30 0C for both tetraspores and carpospores.The spore output in other temperatures were cemparativelylow (Fig. 33) .
4.5.5 PHOTOPERIQQ
Figs. 34 to 40 gives the quantity of spores dischargedfromGelidiel1a_ggg;g§a, Gracilaria ggrticata,.§. edulisand Hypnea musciformig with the effect of different lightand dark cycles on spore shedding at three different lightintensities. Data were obtained at a low light intensityof 500 luxg at a medium light intensity of 2000 lux and ata high light intensity of 4000 lux for all the four redalgae. These light intensities were selected dependingupon the daily spore output observed in these algae(Figs. 26 -_29). In these experiments, planned to studythe combining effects of day length and light energy, peakspore output varied with the duration of light intensity indifferent light and dark regimes.
In=§e1idiella'ggg£g§§ at a low light intensity of 500 lux,tetraspore output increased from O + 25 LD cycle andmaximum sporulation.was observed at 12 : T§'hrs LD cycle.With further increase in light period, though the sporeoutput decreased (Fig. 34) the values obtained were higherthan that at 0 +'§5 LD cycle. At 2000 lux light intensity
Figure No.26 Effect of light intensity on theshedding of tetraspores inGelidiella acerosa.
TETRASPORES / g.fr.w1. / day
04 G3 1
F|GURE.26 GELIDIEL L A ACEROSAXIOOO
96-‘
84-‘
~J N 1
(D O L
J5 m 1
N -I31
R3 1
0 I 1 I 7F’ I0 500 I000 2000 3000 4000
LIGHT INTENSITY (lux)
-"1
Figure No.27 Effect of light intensity on thetetraspore and carpospore outputof Gracilaria corticata.
FIGURI:-$.27. G R ACIL A R I A CORT| C ATAXl000-+ ”
35- o———-0 TETRASPORESA--AcARPosPoREs
0:‘P
N U1 1
20
I54
SPORES /g.fr.w1 / day 2?
0 500 |000”20T00 3000 4000LIGHT INTENSITY (lux)
Figure No.28 Effect of light intensity on theliberation of tetraspores andcarpospores in Gracilaria edulis.
F|GURE.28. GRACIL ARIA EDULI SX I000
40*
04 O J
NO I
23u
SPORES /g.fr. wt. / day
5J
N ‘T
o———o TETRASPORES
A——i\cARP0sP0Rt-:s
A
L
0+4T 7’ I I TI
500 I000 2000 3000 4000LIGHT |NTENSiTY (lux)
Figure No.29 Effect of light intensity on thetetraspore and carpospore outputof Hxgnea muscifdrmis.
F|GURE.29. HYPNEA MUSCIFORMIS
XIOOO 070- O-—-—-O TETRASPORES
Zk——A CARPOSPORES
sod
U1 0 J
-D O ID
SPORES /g. fr. wt./day
N om0 o I l
>
0 1 I I "Ill I I0 500 I000 2000 3000 4000
LIGHT INTENSITY IIUXI
Figure No.30 Effect of temperature on therelease of tetraspores in Gelidiellaac erosa o
F|GURE.30. GELIDIELLA ACE ROS AXIOOO
on ‘B
430) J
TETRASPORES / g.fr.w1.
7? E23 8
I I
0 1 I I I I I-l5 0 5 20 25 30 35 40 45 50
TEMPERATURE (°C)
Figure No.31 Effect of temperature on tetrasporeand carpospore shedding in Gracilariacorticata.
FIGUREL51. GRACILARIA CORTICATAX I000
I6
l4fi
R3 1
51
co 1
SPORES / g.fr.wt. / day
4> as L 1
A
Aw’;
0-——oTETRASP0RES
A——AcARPosPoREs
A \‘L-|'5 V0 20 2'5 3'0 3'5 4'0 415 50
TEMPERATURE (°C )
Figure No.32 Effect of temperature on liberationof tetraspores and carpospores inGracilaria edulis.
spom-:s /g.fr. w1./ day
FIGURE .32. GRAC|LAR|A EDULI SXIOOO32W ,
o——o TetrasporesN (D
I 0
A-——wACorpospores
N J: J
m‘?
S I
I2-+
6- \KO \""""'-—._‘ “..:_“‘-— AI T I I I I I
-IS 0 5 20 25 30 35 40 45 soTEMPERATURE (°C)
F1gure.No.33 Effect of temperature on tetrasporeand carpospore output in Hzgneamusciformis.
F|GU.RE.33. HYPNEA MUSCIFORMISX I000
I00
0——0 Tetraspores
>90- A-——%\CorposporesO
‘O
\, 75; .L.‘
**-.0,60-' . A\6/)LuQ: 45"‘Oo_ Am a30- '\I54 .
0‘ ‘ ’ I 1 I I I-l5 0 5 2025 30 35 40 45 so
TEMPERATURE (°C)
Figure No.3# Combined effects of photoperiod andLight intensity on the tetrasporedischarge in Gelidiella acerosa.
A.m:.3
um.
N_uN_
oo_mmao_.o:a
murw ON.n¢
O‘.
W
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zoom4 469.68Bow80.,
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.3. ooom ollo.:._oo.n_ Tl.
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<m ommoq
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AVG/WfllC]|HO|.LS/S3HOdS V8131
53
the values obtained at 4»: §5 LD cycle were higher hhanin complete darkness (0 + Z5 LD cycle) and peak liberationof spores was higher observed in 12 3'7? LD cycle. Sporeoutput decreased with further increase in the photoperiodat the intensity after 12 : T2 LD cycle. At 4000 luxlight intensity, spore output was high at 4 :'§0 LD cyclethan at complete darkness (0 lux) but spore number wascomparatively higher than that obtained at 8 :'TE LD,12 L : ‘Ti D, 16 L : E (D to 21+ L : 6 D photoregime (Fig. 34)..
In Gracil§r;§_gorticata (Fig. 35) at 500 lux light intensity,slight increase in tetraspore release was seen from 0 +'§Kto 12 :'T§ hrs LD cycle, after which spore output increasedgreatly and highest number of spores was observed at 24 :'0 LDphoto regime. At 2000 lux, peak liberation of tetraspore wasseen at 0 : 25 (complete darkness), whereas in 4 L : §0'Dphoto regime spore liberation was higher than that observedfor other photo regimes (i.e., 8 : TE'LD cycle onwards)shown in (Fig. 35). At 4000 lux light intensity, maximumspore liberation was observed at 4 : 2'6 LD cycle and atother light and dark period spore output decreased (Fig. 35).In the carpospore output of the same species (Fig. 36) at
500 lux, maximum spore liberation was seen at 24 : 0.LDcycle. Spore output increased with the increase in theduration of light intensity. Spore output at 0 light.intensity (complete darkness) was also higher than that
Figure No.35 Combined effects of photoperiodand light intensity on the tetrasporedischarge in Gracilaria corticata.
69:: oo_mmao._oza
butm wuom bum. , w._”N_ murm mm”? w.~.o
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..
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54
observed at 4 ; E0 LD cycle (Fig.36). At 2000 lux lightintensity, spore output increased with the increase in theduration of light intensity and peak liberation.was observedat complete illumination (24 +45 LD cycle), In 4000 lux,maximum spore shedding was observed at 4 : 56 LD cycle. At
8 : lg LD cycle spore output decreased (Fig.,36)..
Fig. 37 shows the carpospore output of Gracilaria edulis.At 500 lux, spore output increased with duration of lightand maximum carpospore release was observed at 20 : 4 and
24 : 5 LD cycle, At 2000 lux, maximum spore release wasseen at 16 : §'LD cycle, Spore output in complete darknesswas higher than that at 4 : £6 and 8’: lg LD cycle (Fig.37).At 4000 lux, light intensity, high spore output was observedat 4 : §5 LD cycle. After 8 : lg LI>cycle. Spore liberationdecreased greatly.
The tetraspore output of Gracilaria_ggulis is given inFig. 38. At 500 lux light intensity, maximum sporedischarge was found at 20 :'4 LD cycle. At photo regimes8pf 16 : 8 and 24 : 5 spore output was more or less of equal
nature (Fig. 38). At 2000 lux light intensity, spore outputwas very high at 8 :'T3 and 12 :'T§ LD cycle. Spore outputat 0 light intensity was higher than that at 4 :'Z0 and16 :‘8 LD cycle. At 20 :'4 and 24 :'U LD cycle, spore
55
liberation was higher than that in complete darkness, 4 :'§5and 16':'§ LD cycle. At 4000 lux light intensity, sporeshedding was high in complete darkness and maximum spores
were observed at 4 : E5 LD cycle, which decreased withincrease in photoperiods. (Fig. 38)
Carpospore liberation of Hypneamusciformis is given inFig, 39, At 500 lux light intensity, spore output increasedwith increase in the duration of light energy; Maximum sporeshedding was seen.at 16 :45 LD cycle. Spore output was alsohigh at 20 2 4 and at complete illumination (24 :'0). At2000 lux light energy, spore output was maximum in dark
(0 light energy) and decreased successively at 4 :‘§5,8:375, 12:73, 2o:'i£an.c124:'5LDcyc1es~. At16‘:-8-LD
cycles carpospore output was as high as observed in 0 lightenergy. At 4000 lux light intensity, maximum spore liberationwas observed in 0 :'§4 LD cycle (0 light intensity). Thevalue obtained at 4 :'§5 LD cycle was less than that obtainedat O 3 E4-LD cycle. But higher than 8 : {E LD cycle. Thespore output then decrease successively in the other LDCycles (Fig. 39).
Figure No.36 Combined Qffects of photoperiodand light intensity on aarposporeoutput in Gracilaria corticata.
69:: oo_mmao»o:a
cm on m. , u. m e o
I A — . % _ 4 .fi q A _ O
noooe
4 4
. loooo
4.V....ooo~_ 8. d4 ..ooom_ %.d.0loooouwm\$ svxJO000nO/noooovmND.400000 0‘
.5. 000¢4|I.Q.
.0000»
.3. ooomollo.2: oom...I...
400005
3: «Solace <_m<.__o<mo .mn daze:
Figure No.37 Combined effects of photoperiodand light intensity on thecar-pospore output in Gracilariaedulis 0
F|GURE..37. GRACILARIA EDULIS (C)- o———-~ 500 lux
IZOOOOF 0-—°2O00lux~ A——A40oouux
lO0OOOJ*'L.
00000“?_3‘ 70000»
‘D\. P‘Q 00000~5 A 4%90- L... 53 aoooot ‘U) L
I110:0 40000?‘ AQ_ AQ) L2 L.0: 30000Q L.L: 20000“ ‘
L
|O0OO‘*r- AQ J I I I 1 I I J I L I 1 [T a0 4 0 I2 I6 20 24
PHOTOPERIOD (hrs.)
Figure No.38 Combined effects of photoperiodand light intensity on the tetrasporeputput in Gracilaria edulis.
A633 ooémaopoza
2 N. 0 ¢ 0
La 14 4. 4 fl A O
‘ O
4
h
I.. 31. ... H0000. VS4 4 d. O1 H3S4 /..6ooo~...o\.
u.m.. ./pD. . .2: 80¢ I A
.5. ooom olld -oooom \
.3. oom fill
CL mflaom <_m<.__o<mo .mm-mm:u:
Figure No.39 Combined effects of photoperiodand light intensity on the carposporeoutput in.Hx2nea mggciformis.
FlGURE'39 HYPNEA MUSCIFORMIS (C)
'---* 500 Iux0""-1'0 2000 IuxA-—A4000|ux/-o
r ~aoooo
4oooq <°
36000 I
/day
25000?
I20000
IIBOOO
CARPOSPORES/g.fr.w1 EO»o I
I39.001')
O O l 1 1 1 1 ‘ 1o 4 a :2 us 20 ' ‘g4PHOTOPERIOD (hr9‘
56
Fig. 40 shows the tetraspore output of Hypnea musciformisat three different light intensities (500, 2000 and 4000 lux).At 500 lux, spore output increased with increase in theduration of light energy from O :'§Z LD cycle and maximum
spore output was seen at continuous illumination (2446 LD
cycle). At 2000 lux light intensity? the value obtained at12 :.?§ and 24»:lO LD cycle showed maximum spore output
than at 0 : ‘£11, 4 _: 2'6, 8 : T6, 16 :3 and 20 :'lI_LD cycles,but W039 51i8ht1Y_ higher than at 0 : EZ LD cycle. At'4000 lux, spore output was high in O +'§Z LD cycle andmaximum spore discharge was seen at 8 : 73 LD cycle. In theother LD cycles (12 : T2, 16} '5, 20 3 '4' and 24 : '5) sporeoutput decreased greatly(Fig.40).
4.6 QERMINATION QF SPOR§§
Data obtained on the percentage frequency of germinating
tetraspores of Gelidiella acerosa and tetraspore andcarpospores of Gracilaria c0rticata,‘§. edulis and Hgpnea.muscifg§g;§ are given in Figures (41 - 44).
GERMINATION OF SPORES IN DIFEERENT MONTHS OF THE YEAR
Monthly means estimated for two years data are plotted toshow the general trend in the germination of spores liberated
in different months of the year. In Gelidiella acerosa thegermination rate of tetraspores varied seasonally from a
Figure No.hO Combined effects of photoperiod andlight intensity on the terasporeoutput in Hxpnea musciformis.
F|GURE.40. HYPNEA MUSCIFORMIS (T)40000
o--- 500 mu
“coop °---0 2000 IuxA——-A 4000 lux
,, 30000»c-u
‘T
‘S 25000:
6x.Eb 20000”u:I:
5' l50001<m.
E;P IOOOO
5060
0 I I 1 J.. 8 I2PHOTOPE R I 00
I6
(hrs,)20 24
57
minimum of 12 % to a maximum of 91 %. Highest rate of
germination (within 24 hours period) was observed inJanuary e February - March and also in July ~ August 83(Fig. 41.1). In tetraspores and carpospores of Gracilariacorticata,;g. ggggig andlfiypggg musciformis, (Fig.41.2-44)germination of spores was observed throughout the year and
highest rate of germination was observed in the months ofOctober, November, December '81, December a January '83 for
tBracilarig corticata and Gracilaria edulis. In Hypneagusciformis highest rate of germination was observed in the_months of August ~ September, 1982, February 1983, July 1983,
for tetraspores and for carpospores from Pudumadam wasobserved in the months of August '82, December '82, January '83,February '83 and October '83, In the carposporic plants(flypneafimuggiformis) collected from Kilakarai, germination
of carpospores was observed highest in the months.of July '82,November '82, and September '83, though germination of sporeswas observed in all the months of the year, which mostlydepended upon the maturity of the spores during liberationand optimum conditions available during and after liberation.
Dividing spores were observed within 24 hours of liberationin Gelidiella acerosa, Gracilaria corticata, Q. edulis, andHypnea musciformig during the two years o£ seasonal studyon spore output.
58
EXPOSURE TO AIR.
Data observed on the germination of tetraspore of Gelidiellaacerosa and tetraspore and carpospore of Gracilaria corticata,‘Q..edulis and HXpnea_musciformis in different experimentsconducted to study the effects of environmental factors aregiven in Figs. 45 -.5h. In Gelidiella acerosa tetrasporesshowed maximum germination in control (d0 minute exposure)(Fig.45A) than the fronds (with fertile stichidia) exposedto 15 minutes exposure and above showed'decreasex ingermination of spores. Spores did not germinate in thefronds exposed to 90 minutes. In Gracilaria corticata,maximum germination within 24 hours was seen in controlin tetraspore while tetrasporic frond exposed to air showeddecrease in germination of tetraspores and there was nogermination in the fronds exposed to 120 minutes (Fig.45B);The carposporic frond exposed to air for 15 minutes showedmore germination than the control. Carpospores alsogerminated.in the fronds exposed to 150 minutes. There wasdecrease in germination of carpospores with increase inexposure after 50 minutes; _There was no germination in thefronds exposed to 2?? hours. In the experiments conductedat room temperature with Qgacilaria edulis, maximumgermination rate was seen in the controls and the germinationrate decreased with increase in the duration of exposure“
Figure No.41 Seasonal variations in the germinationof tetraspores of (1) Gelidiella acerosaAND (2) Gracilaria corticata.
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omssd0om H39
oo_
Figure No.42 Seasonal variations in the germinationof (1) carpospores of Gracilariacorticata (2) tetraspore of Gracilariaedulis.
+540
O
m<4_o<m
m_.Som <_
1 1 4 fl 1 q
<._.<U_._.mOU <_m<.:o<mo
OON
OQ‘
6co
02.o¢om
S380'dSOd8VOS’ 3 8 0d S :10
sauodsvuxa;NOLLVNIIN 3 3 9 as
Figure No.43 Monthly variations in the germinationof (1) carpospores of Gracilaria edulis,(2) tetraspores of Hxpnea musciformis.
FlGURE.43. GRACILARIA EDULIS
.8LA..+Iu
4J38 1M”W .6.
R 4MO 1....._.. 3
C 10.1.3m -n HM. :0 _.l
13
A NE LA 0N ;T.....?_M
L ...J%H .M1
1AJMIF.JlD\|I.
._N..11 2 8P P L[ L L F L F L LL LIL L.L L P L! L
m. w m m o m W W m m o
mumoamoamqu mmmoam<mLmLmmmomm ...o ZO_.r< Z _ 5. m m 0 .\.
Figure No.44 Monthly variations in the gérminationof carpospores of Hzpnea musciformis(1) from Kilakkarai and (ti) fromPudumadam.
zqoqzaosa
_<m$.x<.:x
L
22 mohzomnz 32a»:
.3. -mm:mE
0ON.
9. %9om 33Uom VWall09 NdVo 0180ON dNO0.:3833dow S0Hon 3S
00..
Figure No.45 Effect of desiccation on spore germination:(A) Germination of tetraspores inGelidiella acerosa.(B) Germination of tetraspore and
carpospore in Gracilaria edulis.(B) Germination of tetraspore and
carpospore in Gracilaria edulis.
'1.
‘I. GERMINATING SPORES ‘lo SPORES GERMINKTING TETR ASPORES
F|GURE-45. GERMINATION OF SPORES
. GEUDIELLA ACEROSA304 0——'''° Teflaspores2o~ r
-+
IOT A0 I I I I T T j I §6 us so 445 so 75 90 :05 I20 I354 GRACILARIA CORTICATA
301 o——-0 CarposporesJ 0-——° Tetrospores
B0 * 3 ' I T F 1 I I I fl ‘V r0 I5 30 45 so 75 90 1055120135150 23 3 3’: 4,O0_ GRACILARIA EDULIS
80" 0---0 Carpospores4 0--*0 Tetrospores60‘
404
20-‘. C0 V I I, I V I r 1 féif ‘W I I I IO '9 I - 2 3 4 5 6 7 8
EXPOS RE TO AIR (minutes) (hrs)
Figure No.46 Effect of desiccation on thegermination of spores (tetrasporesand carpospores) in.H1Qnea musciformis.
N A.mL..: Am2::_c.: E4 OH mmamoaxm
e an m «N om: we o~_ mo. cm 9. om 9 on m. o
— _ F\ _ . .4. L r . L .1 pl F O
..o_4' 18nonmmmon_mo.E<o o.||a . now
mmmoamqmpmk ollno
m_sEoh__um:s_ 32¢»: :8
mmmOn_m m0 ZO_._.<Z_S_mm_0 .m¢..mm:oE
°/oS3HOdS €)Ni.LVNlW<‘:l39
59
to air upto 2V2 to 3 hours (Fig.45C). Germination of sporewas not seen in the frond exposed to 3 hours and 392 hours
in tetraspores and carpospores. In the experiments conductedwith Hxpnea_musciformis, maximum spores germinated in thecontrol, and the germination rate of tetraspores andcarpoapores was high in the controls which decreased withthe increase in duration of exposure to air. Tetrasporesdid not germinate in the fronds exposed to 105 minutes andcarpospore germination was not seen in the fronds exposed
to more thar 155 minutes (Fig.-46).
S NITY
Germination of spores tested in different salinities for thefour algal species is given in Figs. 47 - 48. Maximumgermination of tetraspores of Gelidiella acerosa was
observed at 20, 30 to 40 %o salinity. Tetraspores alsogerminated at 50, 60 and 10 %o S, but the intensity waslow; There was no germination at O %o and 70 %o and
above salinity. In the experiments conducted for_gracilariacorticata, gracilariauggglig andlfixpggg musciformis,tetraspores and carpospores did not germinate at O o/oo S.At1O9$o3spores of the 4 algae germinated except the tetraspores ofHxpnea musciformis. Germination rate increased at 20 o/oo
and the highest rate of germination of spores could beseen at 30 o/oo S in.Graci1arga.gorticata, Gracilaria
Figure No.47 Effect of salinity on (1) germinatienof tetraspore in Gelidiella acerosa,(2) germination of ¥e¥raspore In.5raci1ariacorticata (3) germination of carposporen Grac aria corticata (4) germination
of ¥eEraspore n rac aria edulis.
PORES
G
N
'/.GERMINAT'
FIGURE .. 47. GERVMINATION OF SPORES fi~ ~ ROS.ofi , GELIDIELLA ACE A
m -4“'60”0: -4040*Q -4
20-1
m -14 O T V f r Y I 1 I 1.¢8OT 2 GRACILARIA CORTICATA._ * . (T)m
9
U)usa:oa.moa.a:<o
" 4 GRACILARIA EDULIS (T)‘"604mK E240g -4cu 20‘F. -JO -Y 1 1 I T I V T. T0 IO 20 30 40 50 60 TO 80
SALINITY ('/co)
Figure No.#8 Effect of salinity on the following:(1) Germination of carpospore inGracilaria edulis.(2) Germination of tetraspore in
Hzgnea musciformis.(3) Germination of carpospore in
Hxgnea musciformis.
‘lo GERMINATING SPORES
CARPOSPORES TETRASPORES CARPOSPORES
'00 FIGURE..48. GERMINATION OF SPORES.1 1 GRACILARIA EDULIS80- J‘'1 //I ~~'50‘T V \\0 \-1 I \401 /I \.1 ,’ \2o~ ,’ \\.1 I ;\O ¢-*""1' 1 1 1 1 1 \‘-1 1 1" 2 HYPNEA MUSCIFORMIS"1
804"1
60J
404
20+
0 ‘1* 1 1 1 1 r 1 1 1 1. 3 HYPNEA MUSCIFORMIS
30-1
'160* A-1 ’ \\40- ’/ \\’ \-1 / \20-‘ I’ \~‘\I ‘~\0'4 I \ .0 :--=‘'‘'. $7“?' 0 I0 20 30 40 50 60 70 80 90SALINITY ('/co)
60
ggggig andԤ1pggg musciformig, though high rate of tetrasporeand carpospore germination was also seen at 20 o/oo and
40 o/oo S5 Germination_o£ spore was not seen above 70 0/00 Sin Gracilaggg oorticata, 60 o/oo S in.g, edulis and 70 o/oo Sin tetraspore and 60 b/oo S carpospore of Hxpnea musciformig.
TEMPERATQEE
The experiments conducted in different temperatures, for thegermination of spores in the_four red algae has beenrepresented in Figs. 49 - 50. Spores(tetraspores) ofGegidiella acerosa did not germinate in the fronds kept at-15 O; 0:0 C whereas spores germinated at 20 0 C andmaximum rate of germination occurred at 30 O C. Germinationof spores decreased after 35 OC. Sporesdid not germinate
at 45 °C and complete inhibition was seen at 50 OC. Sporegermination tested for the tetraspore and carpospore of_g§acilari§ and Hxpnea for the experiments conducted at
00 010 different temperatures (-15 0, 0 ,55 °, 20 °, 25 ,30 °, 35 °, 40 °, 45 ° & 50 °c) are givenin mgs.49& 50.The tetraspores and carpospores of Gracilaria corticatadid not germinate at -15 QC, 0 . and 5 03; At 20 CCgermination rate increased with maximum rate of germination
was occurring at 30 OC, At 25 ° and 55 °'c also germinationrate was more or less high. .But there was no germination
Figure No.49 Effect of temperature on the :(1) Germination of tetraspores inGelidiella acerosa.(2) Germination of tetraspores inGracilaria corticata.(3) Germination of carpospores inGracilaria corticata.(4) Germination of tetraspores inGracilaria edulis.
GERMIN-ATING SPORES (‘/4
TETRASPORES CARPOSPORES T :1 R A 5 pg 3.; 5
FIGURE - 49. GER MlNAT|ON OF SPORESso
, GELIDIELLA ACEROSAso»
40
0 1% Y 1 1 V D60__ GRLCILAEIA C0'RTICA'TA (T1)2 :40-*
0 9 .. 9 I I 1' ? t I r '80% G-CORTICATA (C).60-‘4°; / ‘K
-0R f04'‘ 7 W fl 1 r r 1 A‘_’T80-1 G.EDUL|S (T)
60-‘
40-‘
20-4
I5 2'0 2‘5 3'0 ‘$15 40 35 soTEMPERATURE (°c)
I 0 Q4
Figure NO 050 Effect of temperature on the following:(1) Germination of carpospores inGracilaria edulis.(2) Germination of tetraspores in
Hzgnea musciformis.
(3) Germination of carpospores inHzgnea musciformis.
(‘Id 1‘,
GERMINATING SPORES
CARPOSPORES TETRASPORES
CARPOSPORES
0 F|GURE-50. GERMINATION OF SPORES.GRACILARIA EDULIS60* ,*. (C)1 , \\,’ \40'" I \\/ \I \201 / K‘I \J \\,_\o-1-0--u=r ‘T T . f . , ‘-r ~4
401 HYPNEA MUSCIFORMIS(T)
2304 ~
20"
I0
0 9 Q T V 1 v _.7 .1 Y T70.. HYPNEA MUSCIFORMISA (C)60-* I \/ \/ \50- II \40.. / X. / \/ \\/30* /, \20- / ,\I I \/ V‘I0‘? // \1 \k0 -J h?’ T I I - I I E r V..‘‘‘? ‘fl?-I5 0 I5 20 25 30 35 40 45 50
TEMPERATURE (°C)
61
at 45 0C in carpospore and at 50 °C for tetraspores. In theexperiments conducted for Gracilaria eaulis spores(tetrasporesand carpospores) did not germinate in the fronds kept at-15 0 and 0 OC. Very few spores germinated at 5 OC andmaximum rate of germination of spores was observed at 25 CC
for both tetraspores and carpospores given in Figs.49 and 50.
There was high rate of germination in the fronds exposed to30 0C; After 35 CC, germination decreased and at 40 DCspores did not germinate and complete inhibition was seenat 45 O and 50 DC. In the experiments conducted for Hvpneamuscifigrjis, tetraspores did not germinate in the fronds
O and 5 OC whereas at 20 OC high germinationkept at e15 °, 0was seen and maximum rate of germination was obtained at
25 DC and 30 OC. Germination of tetraspores decreased after35 0c, at 40 OC spores did not germinate. The carposporesgerminated at 5 OC. Germination of spores rapidly increasedafter 20 CC and maximum germination was observed at 30 °C;
High values were also observed at 25 O and 35 °C and verylow at 40 DC. Spores did not germinate at 45 Oand 50 °C;'
62
LIGHT INTENSITY
The results of the experiments oonduoted at different lightintensities (0, 500, 1000, 2000, 3000 and 4000 lux) for thefour agarophytes to observe the germination rate intetraspores and oarpospores, are given in Figs.51 and 52.The germination rate in tetrasppres of Gelidiella aoerosawas high from 0 to 1000 lux light intensity and maximumrate of germination was observed at 200 lux light energy.Germination rate declined at 3000 lux and spores did notgerminate at #000 lux. (Fig.51.1). s;m11ar1y“1h the "experiments ooEdu©ted‘forvGraéiIaria corticata (Fig.51.2)the values obtained at 500 and 1000 lux were lower thanthat obtained-in dark (0 1ux)} The rate of germination ofspores was high between light intensities of 1000 and 3000lux. In the carpospores the germination observed at 500 luxwas less than that in dark but maximum germination of sporesvv 05
Have observed between 500 and 3000 lux light intensities."In the germination of tetraspores of Gracilaria edulis(Fig.51.&) the rates of germination at 500 lux and 1000 luxwere lower than the value obtained at 0 lux light intesity,Maximum germination.was observed between 1000 lux and 3000
lux light levels. "Germination rate decreased at 3000 lux._The rate of germination in the carpospores obtained is givenin Fig.52.1. At 500 lux and 1000 lux the rates of germination
Figure No.51 Effect of light intensity on the following:(1) Germination of tetraspores in
Gelidiella acerosa.(2) Germination of tetraspores inGracilaria corticata.(3) Germination of carpospores inGracilaria corticata.(4) Germination of tetraspores inGracilaria edulis.
GERMINATING SPORES (O/O)
CARPOSPORES TETRAS PORES
TETRASPORES
F|GURE_51. GER MINATION OF SPORES
80*
60
40-1
20-4
1
GELlD|ELLA ACEROSA "(T)
601
40-+
20-1
GRACILARTA CORTICATTAT
(T)
'1T
a——
T WGRACILARIA CORTICATA
(C)
....——\ \
T “_4
500 I000LI G H T
GRACILAFTIA EDULIS
(T)
3000(lux)
T
2000INTENSITY
400(
Figure No.§2 Effect of light intensity on the following:(1) Germination of carpospore in.Grac11ar1aedulis.
(2) Germination of tetraspore 1n.Hzpneamusciformis.
(3) Germination of carpospore in.HzEneamusoiiormis.
G ERMINATING S PORES (°/o)
CARPOSPORES
CARPOSPORES
IEIRASPORES
F|GURE..52. GERMINATION SPORES
‘ GRACILARIA EDULIS (C)80
A// \\60"‘.\\\ // \\ / \‘\ ______....o’ \40‘-1 ‘v-*’ \\\ \20- ‘~\\\\o f . 0 1 . r \‘:_ HYPNEA MUSCIFORMIS (T)
30- 2
60
40«
200 T Y I I V T__ HYPNEA MUSCIFORMIS (C)
so-4 3,.Q----—''''''—‘X? \.. / \60 Ixz \.2 I \O” \ °40~ \
\ \\20- \\O Y I 1 I * \‘?0 500 I000 2000 3000 4000LIGHT INTENSITY (lux)
63
are observed lower than at 0 lux light level and high sporegermination was seen between 1000 to 3000 lux light level.After 3000 lux, the germination of spores decreased, In
similar mannerrexperiments conducted &3b.H1pnea musciformis(Figs.52,2 & 3): the rate of germination of tetrasporesincreased with the light level and high rate of germinationwas observed between 500 and 3000 lux light energies. Thegermination of carpospores also showed more or less the sametrend (Fig.52,3) i.e. the rate of germination of oarposporasof Hgpngglmusigprmis increased with increase in the levelsof light intensity and high rate of-germination.wasobserved between 500 and 3000 lux. In all these four algaestudied there was no germination at 4000 lux light intensity.
PHOTOPERIOQ
In the experiments conducted to study the combined effects
of selected light intensity with duration of day length onthe germination of spores of the_four red algae studied arerepresented in the Figs.53.1, 53.2 & 53,3).'The tetrasporesofI§g;;gie;£§ acerosa showed high rate of germination withduration in the increase in photoperiod of low light intensity(500_1ux). The values in germination of spores at 4 :‘§5,8 : :3 and 12 : T? to 16—:'§ were low as compared to thefronds in dark (0 light intensity) but in (20 :'H & 24 :;0)_LD cycle higher rates of germination of spores was observed}
64
The fronds that were exposed to 2000 lux light intensityat different photoperiods, high germination was observedat 16 : 3 LD photoregimes, whereas at 20 : E} 24 2 Uand O lux (O : EE) spore germination was higher thanduring other photoperiods. In the fronds exposed to 4000lux light level at different photoperiods, high sporegermination was seen in dark (0 : §E)Au The tetrasporesgerminated,-in the fronds kept,et-4 : E5mLBacyo;e, butwere less than that observed in.O : E4 LD cycle.Germination of spores decreased after 4 : 75 LD cycle andsporesdid not.germinate after 12 :'7E LD cycle (Fig.53.1).
The experiments conducted to study the germination of spores(tetraspore and carpospore) of .§rgg;;aria_gorticata under
different photoperiods are presented in Figs. 53.2 & 53.3.Higher spore germination was observed in dark (0 : E4 LD
cycle) then with the fronds illuminated at 4 : §U & 8 : 36“LD cycle (Fig.53.2). With the increase in the photoperiod.rate of germination of tetraspore increased after 16 :'§ LDcycle and high rate of germinating spores were observed
at 24 :‘5 LD cycle, in the fronds kept at 500 & 2000 1uxplight intensities. Fronds kept at_1ight energy-of 4000 lux,spores germinated in dark (0 light energy) and at 4 :'§5aand8 :‘?6 LD cycle. After 8 :'?g LD cycle germination decreased.
Figure No.53 Combined effects of light intensity andphotoperiod (LD cycle) on the following:(1) Germination of tetraspore inGelidiella acerosa.(2) Germination of tetraspore inGracilaria corticata.(3) Germination of carpospore inGracilaria corticata.
°/.. GERMINATING. sponas
cmpospones T ETRAS Pones
F|GURE-53. GERM|NAT|ON OF SPORES
BOL ‘ GELIDIELLA ACEROSA (T)60‘ 40*
+
20*
O L 4 '1 _ A 580;, 2 GRACILARIA CORTICATA (T)sou ‘
A.. A40L A _ .r 0 ° '20_ A0 1 l 1 3
_ 3 GRACILARIA CORTICATA (C)80- 0-0 500 lux
- A'“13 2000 Iux50L "—‘ 4000 Iux.“ ‘ 50 A ;40. g’ .. ' $ 4,:20» °F V \10 ‘ 1 I 1 I 1 H‘
0224 4220 8I|6 |2:|2 I628 20:4 24:PHOTOPERI OD
Spores did not germinate in the fronds kept under 20 :'Eénd continuous illumination (24 :'5) (Fig.53.2).
Maximum carpospore germination was obtained in the
oystocarpic fronds kept under continuous illumination of500 lux light intensity (Fig.53.3). The values observedin the fronds kept in continuous dark were more or lesshigher than those values obtained in other photoperiods(4 : ‘E6, 8 : T6, 12 : T2) and 16 : 3 LD cycles). withthe light intensity of 2000 lux (medium light intensity),the cystocarpic thalli were tested under differentphotoperiods of 0 + 4 +30, 8 + I‘-IE, 12 + -1.5, 16 + E,20 +‘Z and 24 +‘0 LD cycles. High rate of germinationwas observed at continuous illumination (24 +'U LD cycle)and also at 12 + {E LD cycle). The fronds kept in dark
(0 + 5;) also showed high germination than those under4:+'§5 LD cycle. The cystocarpic thalli kept under highlight intensity (4000 1ux)and different LD cycles astreated above; germination was observed in 0 +‘§Z LD cycle,
4 +.§0 and 8 +'Tg LD cycles, but high rate of germination
occurred at O +'§Z LD cycle and slightyless values wereobtained in 8 +-73 LD cycle (Fig. 53.3). Germinationwas not found at 24 +'U LD photoregime.
Figure No.5¢ Combined effects of light intensityand
(1)
(2)
(3)
(4)
LD cycle on :Germination of tetraspore inGracilaria edulis.Germination of carpospores inGracilaria edulis.Germination of tetraspore inHggnea musciformis.Germination of carpospore inHxpnea musciformig.
GERMQINATING
7. S P 0 R E S
TETRASPORES CARPOSPORES TETRASPORES
CARPOSPORES
FlGURE.54. GERMINATION OF SPORES
80 ‘ GRACILARIA EDULIS (T)L _ :00 ar~**?e0~40» “§f"20r
+0 1 A I L ; _ :0 2 GRACILARIA EDUL|S(C)
e0»
60 - ‘o ° A . ‘T ‘ A ‘40* A _20' o——0500Iux
_ A--A 2000 lux'--* 4000 |ux'0 I A 1 1 A .._ 3 HYPNEA MUSCIFORMIS (T)
60
4o~‘ .4j‘j;§ ‘ ‘ .20
Ar- 'O J 1 J J L 1_ ' HYPNEA MUSCIFORMIS (C)
4;60- #Jw,r- _. 740¢20» i
?r‘O 4__ 1__ L 3 . _0:53 4:20 8:l6 I2~T’é |6:§ 20:4 24:0
PHOf0PERI00
The trend was also similar in'§§gg$;g§;§ edulis forgermination of tetraspores and.carpospores which was seenin dark as well as in continuous illumination of low lightintensities of 500 and 2000 lux and in high light energyof 4000 lux. Spores germinated in dark and the germinationdecreased with increase in the photoperiod of 4000 luxlight intensity. Tetraspores did not germinate aty.20 :‘Eand 24 : 5 LD cycles, while in carpoapores high rate ofgermination was observed at BXTE LD cycle (Fig. 54.2).
lJ1}figyg§;ggsoiiogg;§ also simi1ar“trenfl«wsaobservod.Figs. 54.3 and 54.4 give the trend observed for the
germination of tetraspores and carpospores at threedifferent light intensities (500, 2000 and 4000 lux).Tetraspores and carpospores showed maximum germination
at continuous illumination of low and medium lightintensities but germination of spores observed in darkwere also more than that seen at 4.50 and 8575 to1653 LD pbotoperiods. After 16{§ LD cycle germination
increased. fikhe experiments conducted at high lightintensity of 4000 lux, maximum germination of sporeswas observed in dark (0.54 LD cycle) and the germinationobserved in 4f§0 and 8575 LD cycle, the values of whichwere lower than in.0.§4 LD cycle. Carpospores germinated
upto 12:7§.ph0toperi0d and at 16:§ onwards carpospores
67
did not germinate. Tetraspores germinated upto 20 :‘Z
with a very low rate and at 24 :'U LD Cycle there W35 H0germination (Fig. 54.3)
flode of apore Germination :In Gelidiella acerosa tetraspores germinate according toone of the various germination patterns described byChemin (1937) and Fritsch (1945) as 3Gelidium type’ ofspore germination takes place which resemble the 'Nemaliontype’ in which the protoplasmic contents of the germinatingspore first migrate into a lateral protuberance (Plate 9A)from which the algal thallus later develops. Thisproduction of intermediate multicellular mass of cells fromwhich an erect shoot and a rhizodial system are later
differentiated is the characteristic of the order‘g5lidiales'.
In_§£ggilgr;§:corticata,‘Q._gggl;§ and Hgpnea musciformisbelonging to the order Gigartinales, the mode of germinationof spores %2§L found to be.similar in both carpospores andtetraspores. Disc—like structures were observed.Segmentation takes place in which first a cross-wall isformed dividing the spore into 2 celled stage. lThis isfollowed by a wall perpendicular to the first one dividingit into a 4—celled structure. This with further segmentation
68
by walls perpendicular and paralell to form a multicellulardisc or mass of cells from which an adult thallus develops.This type of germination is described as iDumontia5 type.
4.7 SEASONAL CHANGES IN THE HYDROGRAPHICAL PARAMETERS :
The data collected from October 1981 to June 1983 on theatmospheric temperature, surface sea~water temperature,surface sea—water temperature, salinity, pH and dissolved
oxygen of sea water in the three collection localities.namely Thonithurai, Pudumadam and Kilakarai are presented
in Figs. 55 r 57. During the period of this investigationthe atmospheric temperature varied from 25.5 to 30.4 0C ;
26.0 to 31.5 0C and 26;4 to 32.1 0C, the sea water temperaturefrom 25.8 to 30.7 °c, 26.0 to 30.7 °c, 26.0 to 5o;7 °c,26.4 to 31.9 0C 3 salinity from 28.28 to 35.77 b/oo,V3OI62 to35.59 hfoo and 30.59 to 36.25 o/oo S and dissolved oxygenfrom 3.34 to 6.14 3 3;§5 to 5Q69 and 2.67 to 6.33 ml/1 atThonithurai, Pudumadam and Kilakarai respectively} The
pH of theffigter varied only from 8.0 to 8.7.
Figure No.55 Seasonal variations in the selectedenvironmental parameters at the 5localities of collection.(e.g. Atmospheric temperature, surfaceseawaterutemperature, salinity, dissolvedoxygen, and pH of the seawater in the‘collection localities) at Thonithurai(Station.I).
mzkzoz
.. .2 4 .2 .._ ., o z 3 ., 2 4 2 .,_ o z o
@N b b % _ — P F _ p L _ F » _ P _ _ m.-NI
..x J.’ .\.,n. .\...n/ O fiO m
0N. \/./. 13.. ._.m
/./Z
Z. x [KL Iwm
1./
om. / _-m~Tm.,mmH3. . (mm: clvm
W U
do? roné ‘mm
3U
V _m; . I_mI01®.®
1 .. ..
n NT a rm? N I. m
U3
mm. LB. m -m m
J3
( «J 13$ .m mon; :3 .95 m
«.6 Ia
8.21m» ._u.23-S._m uu<.,E?m 4...: ram 0
.u..u¢..E:.u..:u» u.5:._mo:»< T o 2.\.m
-\_-B zU@>X° I
.2... C._z..:...... I :co:o.m. Zmnxtzoxp .mm-mm:o:
Figure No.56 Seasonal variations in the environmentalparameters at Station II (Pudumadam).
an
mxwzoz
ON 1 0 no.»IN 1 ~ -~.m.3: .. ..¢.m
I I 1
um» um uw.m
r: T I, , ;
.o......au» u.§a.<um uutxam 1...; an . m :3
3.. .._:u» ...:.< 1..l.a
:‘ Xlu..|X I II I
2.... xuo»xoo.IIo
:3 ».:z_..<a oils A
OL¢ r O . -9.»
A HH cote? V
3223.; .wm.mm:o..._
.....mEs~o ma
Figure No.57 Seasonal variation in the environmentalparameters at Station III (Kilakkarai).
(3.) 3Hn.LVH3dW3.L
m I F 2 O 2
now. _ - mom. _ _mm_
3 I 4 E . u. _. 0 Z O m 4 1 3 S < 2 ... 1 0 Z O
QN r P . p LI p p L » » Fit _ IF L _ ? p . ? » h 051 O 30rm; 5...S. L?»
L F
.T rm.»mm; 8.. saw» ¢u»<;-<mm mafia» 4...... umn r o rm.»
8.. u¢=»<¢u..au» u:_u:..oa»< I
L It XI..I.X I. T T
.2... zuorxo o..l.o:2 »tz_..<» I
9.: re... re. ro.m..\.m Seuo. 1..
nHHH 3:23 _<m<xx<._:. .$-um:oE
69
Ct , ER- 3DlS_ ONTIn the present study, data were collected on fruitingbehaviour and sporulation of four economically important redalgae growing along the Mandapam coast namely Gelidiella
acerosa, Gracilaria orticata,‘g, edulis and Hypneapmusciformfis.Experiments were conducted with these algae under laboratoryconditions to find_out the variations in spore production indifferent months of the year and to know the diurnalperiodicity in spore shedding and environmental factorsaffecting the spore output.
As reported earlier from Mandapam area for Gelidiella acerosa(UmamaheswaraRao, 1973 a; Thomas et al., 1975 a; Rama Rao
and Subbaramaiah, 1979; Subba Rao, 1979 and Chennubhotla;g§;§;a,1979); Graoilaria corticata (Umamaheswara Rao, 1972 b and
' and §.edulis Cumamaheswara Rao 1973 b andChennubhot1a_§§_§;.; 1979);Ain the present investigation alsoChennubhofla et a1 1979)populations'bf"these red algae and also Hypnea musciformiswere observed throughout the year in the vicinity of Mandapam.
FRUITING BEHAVIOUR
At Veraval the tetrasporophytio plants of Gelidiella acerosawere seen in the population during April -May and October November (Sreenivasa Rao, 1974). The tetrasporophytes in thepopulation of Gelidiella acerosa growing at Rameswaram remainedcontinuously for 7 to 9 months from February to October and
70
in general maximum number of plants with stichidia occirredin the population after the two peak growth seasons in anyear (Umamaheswanl Rao, 1973 a). In the Qglidiella acemasaplants growing at Pudumadam, tetrasporic plants occurredin all the months of the year except in July. Minimumnumber of stidhidia with mature tetrasporangia was found inAugust and September.
Similarly the number of stichidia per plant was low againin February and March. In other months of the year fertileramulii were more, indicating that peak reproductiveactivity occurred between April and June and betweenOctober and December (Umamaheswara Rao, 1974). At Kilakarai
plants of Gelidiella acerosa showed peak reproductive periodof tetrasporic plants in.June (Thomas_g§_§;., 1975 b),whereas in Gelidiella acerosa growing at Rameswaram thenumber of stichidia per plant showed a maximum in.Januaryand the maximum percentage of tetrasporic plants wasobserved in June. A second lower maximum in the number ofstichidia per plant was observed in August. The reproductivecapacity of the population which combined both these
features showed two maxima, one in January and another inJune. The tetrasporic plants occurred only from August
to January in a year (Rama Rao‘g§.g;., 1976). As reportedin other geographical areas (Chihara and Kamura, 1963)
71
sexual plants were not encountered in these studies . Butin the present study tetrasporic plants were found throughoutthe year in the populations of §elidiella'acerosa growingat Kilakkarai as previously observed at Pudumadam by
Umamaheswara Rao (1974). Cystccarpic plants in Gelidiella
acerosg were not encountered in the present investigationas reported from India by Sreenivasa Rao; (19$9) andUmamaheswara Rac (1973 a) and in other geographical areasby Chihara and Kamura (1963).
In_Gracilar;§_corticata growing at Veravalg tetrasporicplants occurred almost throughout the year while sefiualplants occurred seasonally (Oza, 1979). But in Gracilariacorticata growing at Mandapam (Umamaheswara Rao, 1976) andVisakhapatnam (Subba Rangaiah, 1978) both tetrasporic andcystocarpic plants occurred in all months of the year.Similarly asexual and female plants were observed throughoutthe year in the population of Gracilaria cortipata occurring‘at Pudumadam“
In Gracilarig edulis growing at Rameswaram, tetrasporophytes
occurred for eleven months in an year and cystocarpic plantswere seen only in the month of January(Umamaheswara[Rao,1973 b)But in Gracilarig edulis growing at Thonithurai, tetrasporicand cystocarpic plants occurred in the population in allmonths of the year. JAt Visakhapatnam coast tetrasporophytes
72
and oérposporophytes in the population of Hypnea musciformisoccurred in all the months of the year (Subba Rangaiha, 1978).Though the tetrasporio plants were observed throughout the
year in Hypnea musoiformis growing at Pudumadam, cystocarpicplants were seen only for some months in the population atPudumadam and Kilakarai. ‘This agrees with the findings of
Rama Rao (1977 b) for Hzpnea musoiformis growing at Pudumadam,cystooarpio plants were seen only for some months in thepopulation at Pudumadam and Kilakarai. This agrees with thefindings of Rama Rao (1977 b) for Hypnea;musCiformis growingat Veraval and fiypnea valentiae growing at Pamban andKrusadai Island.
§pore Shedding :
Maximum and minimum values obtained on spore productionduring the entire period of this study are given below toshow the range in the quantity of spores liberated inGelidiella ggggggg, Gracilaria corticatg, §,lggg;;§ andHxpnea.Qusci£prmi§.
73
Table (9)
Maximum and Minimum Values of Spore Output :
(expressed as g. fr. wt. per month)
Algae Esfaf aasiéa %‘:“:%3%?9a%;:% 3'Minimum. Maximum Minimum Maximum
Celidiella adasagg 5,744 5,89,793 9- 7Gracilaria_ggrticata 8,181 92,277 8,086 4,87,178g. eduiig 2,155 -1,o9,565 28,652 13,27,853Hzpnea musciforggg 2,662 5,04,953 2,814 6,46,385221.‘.-L1‘ . '"
Studies that have been conducted on sporulation of somered algae growing at Mandapam area and other localities ofIndian coast are discussed below :
In Gelidiella acerosa a maximum output of 10,000 tetrasporesper plant was observed (Sreenivasa Rao, 1969); The tetrasporeproduction varied from about 5000 to 10,000 spores/g. freshwt./day in the plants collected from Pudumadam (Umamaheswara
Rao, 1974 a). The tetraspore shedding in Gracilaria corticataoccurring at Mandapam ranged from 84,000 to 3,98,000 spores/g.fresh wt./day and carpospores from 1183 to 2374 spores per
cystocarp per day (Umamaheswara Rao 1976). A maximumcarpospore liberation of 8,66,700 spores/plant was found
74
by Mohan Joseph and Krishnamurthy (1977) in Gracilaria
cortioata collected from.Mandapam.
In Gelidium pusillum and Pterocladia heteroplatos growing
at Visakhapatnam coast, tetraspore output ranged from.1,149 to 10,78,505 and carpospores from 1,176 to 6,99,943in Gelidium_pussilum and 1,427 -1 7,94,055 tetraspores39,966 - 1,67,04O carpospores/g. fresh wt./day in§terocladia_heteroplatos (Kaliaperumal 1979).
In-Graoilar;§;corticata occurring at Mandapam, tetrasporeshedding range: from 84,000 to 3,98,000 spores/g. fresh wt./day and carpospores from_1,183 to 2,374 spores/cystocarp/day(Dmamaheswara Rao, 1936). A maximum carpospore liberation
of 8,66,70O spores/plant was found by Mohan Joseph and
Krishnamurthy (1977) in.§. corticata collected fromMandapam, The carpospore output per plant varied from
6,919 to 6,49,873 from plants of Gracilaria edulis collectedfrom Krusadai Island near Mandapam (Rama Rao and Thomas,
1974).
In Gracilarga millardetgg, Krishnamurthy (1967) recordedmaximum output of 68,500 tetraspores and 42,782 carpospores
per plant. Oza and Krishnamurthy (1968) observed a maximumliberation of 70,000 oarpospores per plant in Gracilaria_yerrucosa growing at Kuda near Bhavanagar in Gujarat. In
75
Hzpnea valentiae £rom1Mandapam maximum number of 3,14,9#4
tetraspores and 7,01,607 oarpospores were released
(Rama Rao,~1979)3 Subba Rangaian (7978) collected data on-etetraspore and oarpospore output: in the four members ofGigartinales namelylggggglggig oorticata,;§. textorii,_§raoilariopsis.gfigestedtgg and Hxpnea musoiformis growingat Visakhapatnam coast and maximum spore output was found
on first day. The tetraspore output ranged from 13,665 to12,30,380 spores/g. fresh wt./day and oarpospore outputvaried from 280 to 6807 spores/oystooarp/day among the four
red algae. In.Gelidiopsl§_Xar}abi1is growing at Visakhapatnamdoast shedding of tetraspores from stiohidia was observedfor 3-4 days with maximum discharge of_spores on the firstday (Kaliaperumal and Umamaheswara Rao, 1982). The quantityof spores liberated on the first day varied from a low valueof 20 spores to a maximum of 2,60,94O spores per gram freshweight of the plant.
The oarpospore output per plant varied from 6919 to
6,49,873 in Graoilarig edulis collected from KrusadaiIsland near Mandapam (Rama Rao and Thomas, 1974).
76
The present estimation given above in Table (9) are basedon total number of spores released per month. The abovefigures reveal that the spore shedding capacity is veryhigh in the four algae studied. The values obtained fortetraspore output in Gelidiella acerosa in the presentinvestifation are higher than that obtained for;Q. aoerosagrowing in the tidalpools at Veraval (Sreenivasa Rao,1969).The tetraspore production varied from about 5000 to 10,000spores/g fresh wt/day at Pudumadam (Umamaheswara Rao,1974 a).
The spore output estimated in the four red algae can becompared with the values reported for Gelidigm amansii(Suto, 1959 b), Gelidiumppussilum and Pterocladia heteroplatqs(Kaliaperumal, 1979), Graoilaria oorticata (Umamaheswarafiao, 1976; Mohan Joseph and Krishnamurthy, 1977 and SubbaRangaiah, 1978),Ԥ. edulis (Rama Rao and Thomas, 1974), Q}
millardetii (Krishnamurthy, 1967),.§. textorii (SubbaRangaiah, 1978),.§. verrugosa (0za and Krishnamurthy, 1958),Egpnea valentiae (Rama Rao, 1979), Hxpneaggusoiformis and
Gracilariopsig sjoestedtii (Subba Rangaiah, 1978) and4§elidiopsis_yariabilis (Kaliaperumal and UmamaheswaraR30 9 ‘o
In general, maximum shedding of spores was seen on the firstday in the four red algae (Tables 1-8) and it agrees withthe results obtained for Gelidium amansii (Suto, 1950 a),
21-.uJ1..J.2 (Mm: 1950) y §.1:.e9.i,ler_i.e edulis(Rama Rao and Thomas. 19?fl},’§,pggrticat§ CMohan Joseph
and Krishnamurthy-, 1977 and Suhba Rangaiah, 1978), EC.‘
gegtorii, Ggaeilarioggis sjoestedtii and Hxpgea musciformgs(Subba Rangaiah, 1978) and oelidium p}_1wsil"ILum, g_t;roc1gd;_g
heteroplagos and Gelidioggispggriabilig (Kaliaperumal, 1979‘and Kaliaperumnl and Umamalmswra Rao, 1382). Rhythmicliberation of Qarpospor¢$ with peaks at intervals of 4-5days was reported in.Graoilaria‘gorti§g§g.(Mohan Josephand Krishnamurthy, 1977), G, edulis (Rama Rao and Thomas,
1974) and Q, yerruoosg (Qua and Krishnamurthy. 1968).Similar trend in the diaohergq or oarpospores was observedin the present study on,Greo1;aria.gprtigat§,_Q..ggg;;§ andHxpnea ggsciformis (Tables 5-8 & Fig. 2.).
Seasonal Changes in Spqrergptpgt:seasonality in the liberation of tetraepores was foundin Gelidie;;§.ggg§g§§ growing in the tidla pools atVeraval (Sreenivasa Rao, 1971 a and 1974). In qelidiellaacerosa growing in the tide pools at Veraval, the liberation of tetraspores oommenced by the end of April and theShedding of spores in.maximum number of plants was over
by the end of May. During October-November there was asecond cycle of formation and subsequent liberation oftetraspores. The production of tetraspores during this
78
period of reproductive activity is less compared with thatin ApriléMay. Moreover the spore production season inOctober-November was shorter than that in ApriléMay. Theshedding period extended only for 25 to 30 days in eachfruiting season (Sreenivasa Rao 1971 a and 1974). Butin the present study on Gelidiellg ggggggg growing atKilakarai shedding of tetraspores was observed throughout the year as reported for Gelidie;;§_ggg§ggg growingat Pudumadam (Umamaheswara Rao, 197h). In Gelidiella
acerqga growing at Pudumadam shedding of tetrasporeswas observed during the entire fruiting period with peakspore output during May—June and November—December. The
spore output was minimum during the two vegetative growthphases of this algae and though the number of stichidiaper plant was more, the spore production was less inApril and October. This may be due to the occurrence ofimmature tetrasporangia immediately after vegetativegrowth periods (Umamaheswara Rao, 1974). Spore shedding
was found in all months of the year in‘§g;;g;gg‘pg§;;lggandmgtgggglggig heteroplatos growing at Visakhapatnam coastwithout any seasonal periodicity in the quantity of tetraqspores and carpospores liberated (Kaliaperumal, 1979). Inthe earlier studies on_§g;;gigl;g_ggggg§g,seasonal vari_ation in the number of stichidia and peak discharge oftetrasporos ht particular periods of the year was reported
79
by Sreenivasa Rao, (1971 a and 197h); Umamaheswara Rao,
(1974) and Rama Rao g§..g;.,(1976). In the presentinvestigation, though spore shedding wastseen in allmonths of the year, in.§g;;gi§;;g_§gg£3§§ seasonalveriations were not found (Fig. 38) and in this aspectit agrees with members of Gelidiales namely Qelidiggml.-°.-.».:;L_1.J.+.L_1.m.. and Pt e_r.:_<>a1adis ;'1.s.ia<2£;9_12.ls.’§,a§ gorwins at
Visakhapatnam coast (Kaliaperumal, 1979). Similarlythere was no seasonality in the production of stichidiain s2s.I;0_§s (Fiso 31*»)
Data on seasonal changes in the shedding of tetraspores
and carpospores of Gracilaria corticata from.Mandapam werecollected by Umamaheswara Rao (1976); Shedding of tetraspores and carpospores was observed in all months of theyear with marked variations in the spore output. Thetetraspore output was maximum in.March and April andanother small peak in NovembereDecember with an increase
in spore output from September. The carpospore productionwas maximum in March and again in September and October.
The seasonal periodicity in carpospore output was somewhat different from that of tetraspores. The second peakin carpospore shedding was observed two months earlier andhigh values were also obtained in January and June. Mohan-Joseph and Krishnamurthy (1977) observed seasonal changes
in the shedding of carpospores with maximum output on the
first day and rhythmic liberation of carpospores was alsofound in this species collected from.Mandapam. The tetra~
spore and carpospore output varied seasonally in Gracilariacorticata and gypgga musgiformig of Visakhapatnam coast,with peak shedding of spores in the periods betweenDecember and February/March and from August to October
each year (Subba Rangaiah, 1978). Two peak periods ofsporulation were observed in}g§gg;lgr;g.gggt;gg£a occurringon the coast of Veraval. Laboratory experiments wereconducted which showed the tetraspore shedding from April
and August and carpospores shedding from November toDecember (Oza, 1979). Studies on the seasonal rhythm in
the shedding of tetraspores and carpospores in fiypgggyalentiae from.Mandapam were carried out by Rama Rao (1979).
Maximum number of tetraspores were released in Februaayand carpospores in October. But similar trend was notobserved in the liberation of tetraspores and carposporesin Gracilaria corticata_ growing at Pudumadam (Figs. 3C
and 3D) in the present study.
Seasonal variations in the liberation of carposporeswith peak activity at particular periods of the year wasreported in.E£§§ilag;§ edulis collected from.KrusadaiIsland (Rama Rao and Thomas, 1974). But in Gracilaria
81
edulis collected from Thonithurai seasonal variations were
not found in the release of tetraspores and carpospores(Figs. 4A.and 4B). The duration of shedding of carposporesin Gracilaria edulis growing at Rameswaram increased from3-4 days in February to 30 days in August and decreased fromSeptember to January; The daily periodicity was alwaysmaximum-in the first day and decreased gradually but showeda second maximum after fourth or fifth day in the months ofJuly-August where the number of days of sheddinfi“was sprolonged. The total spore output per plant showed peakvalues in July and August which gradually decreased byJanuary. Higher values of spore output were seen again inFebruaryémarch while in April and.May a total lack ofspores was noticed (Rama Rao gnd Thomas, 1974). Seasonalvariations in the liberation of spores with peak aptivityat particular periods of the year are reported inGracilaria verrucosa (Jones 1959 a and Oza and.Krishnamurthy,
1968) and gelidiopsis variabi1is(Kaliaperumal andUmamaheswara Rao, 1982). Rhythmic_1iherotion of carposporeswith peaks at intervals of 4-5 days were also recorded inGracilaria verrucosa by Oza and Krishnamurthy (1968).Similarly as observed in the present investigation, therewas no seasonal periodicity in the shedding of tetrasporesand carpospores of Hypnea musciforis collected from
82
Pudumadam and Kilakarai (Figs. 4C and 4D) while it wasobserved in Hypgea'musciformis from Visakhapatnam (Subba
Rangaiah, 1979), and Hypneavalentiae from Mandapam(Rama Rao, 1979).
‘Qiurnalgperiodicity in Spore Shedding :Diurnal periodicity was observed in some members ofGelidales and other red algae with maximum shedding ofspores at a particular time in a day. Diurnal periodicityin the liberation of spores with maximum output at aparticular time in a day was not found in Iridophycuspgurnucopiae (Fukuhara, 1957). But periodicity in theliberation of spores was observed in members of Gelidialesand other red algae (Suto, 1950 a and b; Katada_g§‘g;.,
1953; Katada, 1955; Umamaheswara Rao, 1974 a; Kaliaperumal,1979 and Ngan and Price, 1983) and Gigartinales (Matsui,1969; Umamaheswara Rao, 1976; Subba Rangaiah, 1978;_Kaliaperumal, 1979: Rama Rao, 1979 & Ngan and Price, 1983).Matsui (1969) observed the release of maximum number of
tetraspores and carpospores from evening to mid-night inGloiopeltis tenax and in the early morning hours inGloiopeltis furcata. Umamaheswara Rao (1974 a) reported
-the periodicity in ‘Gelidiella acerosa with peak sporeoutput in the day-time between 2 PM and 6 PM.
83
In Gelidiella‘ggg£g§g growing at Kilakarai also diurnalperiodicity with peak liberation of tetraspores in theafternoon from 2 P.M to 6 P.M. was found in the presentstudy. It confirms with the observation made by UmamaheswaraRao (1974 a) on the diurnal periodicity in the output oftetraspores in Gelidiella‘ggg£g§g growing at Pudumadam.Seasonal variations on the diurnal periodicity of sporeoutput was observed in Gelidiales members such as Gelidium
amansii (Katada_gt.a;., 1953) and Gelidium pusillum andPterocladiglheteroplatos (Kaliaperumal, 1979). But therewas no change in the diurnal periodicity surges obtained fordifferent months from October 1981 to September 1983 inGelidiella acerosa (Figs. 5 and 6) and it is in confirmitywith the results obtained for Gracilaria_ggrticatg, Q,textorii, Gracilariopsis_§joestedtii and_§ypggg musciformisgrowing along Visakhapatnam coast (Subba Rangaiah, 1978).
Suto (1950 b) and Katada _e_t _a_I_L_., (1953) found the shedding ofspores in Gelidium amansii daily in the afternoon.
In gracilaria corticata maximum spore output was observedduring night time between 10 PM and 6 AM. (Umamaheswara
Rao, 1976). Subba Rangaiah (1978) observed peak shedding
of tetraspores and carpospores in ggacilaria corticata andggacilariopsis gjoestedtii and tetraspores of Hypneagusciformis during night time between 2 AM and 6 AM.
In ggacilaria textorii maximum liberation of tetraspores aswell as carpospores was seen in the afternoon between 2 PMand 6 PM. Data collected continuously for 2 days withtetrasporophytes and 3 days with cystocarpic plants indicatedthe ocourrence.of similar diurnal periodicity in theliberation of tetraspores and carpospores of the above algae.
Diurnal periodicity with a prominent peak in the liberationof spores was observed in Gelidium_pusillum and Gelidiopsis
_variabilis and a definite peak in spore shedding was notseen in Pterocladia heterqplatos (Kaliaperumal, 1979).There was no difference in the time of peak shedding of
carpospores and tetraspores in Gelidium_pusillum and
_Eterocladia heteroplatos. In Geligium_ppsillum peak outputof spores was observed during night time between 6 PM and
10 PM or 10 PM and 2 AM. In Gelidiopsis variabilis peakshedding of tetraspores was found during the day time from6 AM to 10 AM or from 10 AM to 2 PM. Rama Rao (1979)
observed a definite diurnal periodicity in the shedding ofcarpospores in.fiypnea valentiae. Ngan and Price (1983)recently collected data on the periodicity of carposporeand tetraspore discharge in 14 ded algal taxa from thevicinity of Townswille region, Queensland, Australia undera variety of laboratory conditions at hourly or bi-hourlyintervals over periods of 24 hrs or in some cases 48 hours.
85
In the diurnal periodicity of tetraspores discharge based onhourly records of spore output, the spore liberation.wasobserved from 04.00 hrs to 10.00 hrs with peak at 06.00 hrs
in Gracilaria edulis. The time of maximum output oftetraspores and carpospores in Graci1aria_§h9dotrichaandyflypggg cervicornis and tetraspores in Gracilaria textorii,‘Q. verrucosa, Hypnea pannosa and‘§. valentiae was found inthe morning between 05400 and 09.00 hours. The aboveobservations is in confirmation with the data obtained inthe present study for the three members of the order
Gigartinales {Graoilaria corticata, Q. edulis and HypneaEyusciformifi).
Differences in the time of peak shedding between tetrasporesand carpospores was observed by some workers. Katada gt 3;.
(1953) found that the shedding time of carpospores in Gelidium.§gggg;; was always earlier (3-4 hours) and shorter thantetraspores. Subba Rangaiah (1978) observed maximum
shedding of carpospores in Hypnea_musciformis 4 hoursearlier (10 PM to 2 AM) than peak tetraspore output. Thedata collected in the present study on Hypnea musciformisagrees with that of Subba Rangaiah's (1978) observations.But there was no difference in the time of peak tetrasporeand carpospore shedding in_gloiopelti§ tenax and Gloigpeltis
furcata (Matsui, 1969),‘0racilariapggrticatg (Umamaheswara
Rao, 1976} and Subba Rangaiah, 1978), Gracilaria textorii
and Gracilariopsis sjoestedtii (Subba Rangaiah, 1978),Gelidium_pusillum and Eterocladia hetgroplatos (Kaliaperumal,1979).
Seasonal variations in the diurnal periodicity of spore outputwere also investigated. Katada'gt.§;. (1953) showed thatthe time of peak shedding of tetraspores and carpospores inGelidiym‘gggg§;; becomes gradually earlier (between 12.00 and14.00 hours) in a day between June and September and itoccurred later-(after 14.00 hours) in a day during Octoberand November. Similarly seasonal changes in the diurnal
periodicity of tetraspore output in Gelidiug pusillum andQfilidiopsis variabilis was observed by Kaliaperumal (1979).In gglidium ppsillum peak output of spores was observedbetween 6 PM and 10 PM from March to November and four hours
later between 10 PM and 2 AM from December to February.
In Gelidiopsis variabilis peak shedding of tetraspores wasfound between_6 AM and 10 AM from April to November andbetween 10 AM and 2 PM from December to March. But in
ggragilaria corticata,‘§. textorii, Gracilariopsis_§1oestedtiiand Hxpnea musciformis the diurnal periodicity of tetrasporesand carpospores did not vary in different months or season ofthe year (Subba Rangaiah, 1978).
Though a definite peak at one period of the day as observedin the present study was not seen in the shedding of
tetraspores»and"carpospores in Graoilarig Qorticata, Q, edulisand Hypnea musciformis, maximum liberation of sporesoccurred in these Species between 10 PM and 2 PM (Figs. 7;to 17).This period coincides with the times at which maximum spore
output was observed for Gracilarig corticata and Hypneamusciformis growing in India (Umamaheswara Rao, 1976 and Subba
Rangaiah, 1978) and Gracilaria edulis and other species ofpggacilaria and Hypnea such as Gracilaria rhodotricha,.§.textorii, Q. verrucosa, Hypnea cervicornis,4§.ppannosa and‘E. valentiae also growing in the vicinity of Townsville region,Queensland, Australia (Ngan and Price, 1983).
EFFECTS_OF ENVIRONMENTAL FACTORS ON SPORE SHEDDING
For a successful cultivation of spores, we need to know thedifferent factors affecting spore output. Effects ofenvironmental factors like desiccation, salinity, light(intensity and photoperiod) and temperature on spore sheddinghave been reported by few workers. Sea water temperature
and the above conditions are considered as important factorsfor controlling the growth and reproduction of intertidalalgal population. Detailed studies on these ecophysiologicalaspects have not been made on the intertidal algae growingalong the Indian shores particularly Mandapam coast. Thescanty information available on spore output indicates thatthe response to various factors of the environment variesin different algae.
88
EXPOSURE TO AIR AND DESICCATION
Katada (1955) reported that desiccation in the shade has noinducing effect upon the shedding of tetraspores in Gelidiumamansii and in these experiments the time of shedding was
extended within about half a day. On the contrary, Matsui(1969), reported accelerating effect in Gloiopeltis species.Accelerating effect in liberation of spore was observed inQl0igpeltiS‘£gg§§ and Gloiopeltis furcata when the frondswere exposed to air.
In Gloiopeltis tenax fronds were exposed to air for 2 ~ 6 hourswhich hastened the liberation of spores and majority of sporeswere liberated within a short time, even 10 hours before thepeak time of daily spore liberation. In Gloiopeltis.£u§g§£§when exposed fronds were reimmersed around the time of peak
daily liberation, a large quantity of spores were liberatedimmediately after reimmersion. Even if the liberation did notoccur immediately after immersion, the exposure affected thetime of subsequent liberation. In both species the number ofspores liberated immediately after immersion gradually decreasedif the exposure was prolonged beyond 6 hours.
In gelidiella acerosa (Sreenivasa Rec, 1971 a) drying oftetrasporic plants in shade had no effect on spore output.The spore shedding decreased in the fronds exposed for 1~2hours and prolonged drying of fronds for 4 to 12 hoursinjured the plants,
89
In Gracilaria cortiggta (Umamaheswara Rao, 1976), tetrasporeoutput decreased markedly in thalli exposed for 1 hour andspore shedding was not seen in fronds exposed for a periodof 2 to 6 hours. Carpospores were not at all liberated fromcystocarps exposed even for 1 hour to air at room temperature.In the experiments conducted with tetrasporic thalli of
Gracilaria corticata,flgracilaria textorii,_gracilariopsissjoestedtii and Hypnea musciformis exposing upto 105 minutesto air in the room with 15 minutes intervals, the tetrasporesoutput decreased in all plants with increase in the durationof exposure of fronds and maximum output was seen under
continuously submerged conditions (Subba Rangaiah, 1978 andUmamaheswara Rao and Subba Rangaiah, 1980).
.In the desiccation experiments conducted with tetrasporophytes
sof_§elidiumppusillum, Pterocladia heteroplatos and Gelidiopsislygriabilis growing at Visakhapatnam coast, maximum tetrasporeAoutput was obtained from the thalli kept under submergedconditions. The above observations agree; with the presentstudy made in_§elidiella acerosa, Gracilarig_9orticata,.§.edulis and Hypnea musciformis.
According to to Umamaheswara Rao and Sanjeeva Reddy, (1982),
maximum tetraspore shedding in Dictyotg dichotoma was seen
control experiments where the fronds were not exposed to air.The tetraspore output decreased even in fronds exposed to airfor 15 minutes. with increase in the duration of exposure
90
to air, the spore output declined rapidly and total inhibitionof spore shedding was found in fronds exposed for 120 minutes.But in the fronds of Gelidiella acerosa, Gracilaria_gorticata,‘Q. edulis and‘fl1pggg musciformis exposed to air, spore outputdeclined than in the fronds kept continuously under submergedcondition (Figs. 18-21) indicating that exposure of frondsduring low tides adversely affects spore liberation. Submergedcondition was found favourable for maximum spore release inthese four algae. These findings agree with the resultsobtained ior.gg;;g;gg gggngii (Katada, 1955), §:;idie1lgacerosa (Sreenivasa Rao, 1971 a), Gracilarialportioata(Umamaheswara Rao, 1976; Umamaheswara Rao and Subba Rangaiah,
1980; Subba Rangaiah, 1978), Hypnea musciformis, Gracilariatextorii and Gracilariopsis gjgestedtii (Subba Rangaiah, 1978;and Umamaheswara Rao and Subba Rangaiah, 1980), Gelidium
_pusil1um, Pterocladia heteroplatos and Gelidiopsis vgriabilis(Umamaheswara Rao and Kaliaperumal, 1983) and_Qictyota dichotoma
(Umamaheswara Rao and Sanjeeva Reddy, 1982). The maximum
tetraspore shedding in Dictyota dichotoma was seen in controlexperiments where the fronds were not exposed to air. Thetetraspore output decreased even in the fronds exposed toair for 15 minutes. With increase in the desiccation ofexposure to air the spore shedding was found in frondsexposed to 120 minutes.
91
SALINITY
Any significant variation was not observed in the shedding ofspores between 17 and 52%osalinity in §;oiopelti§_tenax andggoiopeltis furcata by Matsui, (1969). More number of sporeswere liberated in 15.6 chlorinity (28.19 %o salinity) and thespore output decreased at 11.0 (19.89 %o salinity) and 8.2ohlorinity (14.83 %o salinity). At 5.4 chlorinity (9.78 %osalinity) the number of spores liberated wase very low.Matsui (1969) found that the tetraspore liberation was notsignificantly altered in salinities between 17 %o and 42 %oin Gloiopeltis tenax and Gloiopeltis_furcata. At salinitiesbelow 12 %o and above 60 %o liberation of spores was delayedand the number of spores also decreased in these two plants.Monospore release was found in Acrpchaetium endophytium'byWhite and Boney (1969) in the media of 19.2 %o --49 %o
salinity, while plants died in 6.4, 12.8 %o S and in salinitiesabove 49 %o. Normal production of zoospores or gametes was
found in Ulva fasciata by Mohsen‘g§_g;., (1972) in thesalinity range between 20 %o and 35 %o. Higher salinitiesfrom 35 to 45 %o enhanced both the formation of swarmers and
their discharge. On the other hand salinities below 20 %oretarded both the maturation and discharge of swarmers andbelow 15 %o no dehiscence took place.
92
The spore output varied in experiments conducted by Subba
Rangaiah g; g;., (1975), Subba Rangaiah (1978) andUmamaheswara Rao and Subba Rangaiah (1980) in different
salinities ranging from 10-60 %o and the optimum rangeobserved for peak shedding of spores was 30-40 %o S. in
Grapilaria cortioata, grggilaria textorii and Graoilariopsisgjoestedtii and 20-30 %o in Hypnea musoiformis. UmamaheswaraRao and Kaliaperumal (1983) conducted spore output experiments
with tetrasporic thalli of Gelidiumlpggglggg, Pterocladiaheterdplatos and Gelidiopsis variabilis in different salinitiesranging from 0 to 70 %o. In Gelidium pusillum and Pterocladiaheteroplatos, spore shedding was found from 10 to 60 %o withpeak discharge at 30 %o salinity.
In gglidiopsig variabilis spore shedding was seen only in threesalinities (20, 30 and 40 %o) with lowest value at 20 %o s.Sa1inities«<20 %o completely inhibited spore shedding and at40 %o salinity the spore output was more than in'gglg;gm_pusillum_and‘gggggpladia heteroplat s Of the eight differentsalinities (0-70 %o salinity) tested with'Q;g3yg£g dichotomaby Umamaheswara Rao and Sanjeeva Reddy (1982), spore output
was observed in salinities ranging from 10 %o to 60 %o;From 10 %o salinity onwards the spore output increased andmaximum shedding of tetraspores was observed at 30 %o salinity.The spore output declined from 40 %o salinity and it wasminimum at 60 %o salinity. But as observed in the present
93
study salinity of sea water influenced sporulation inGelig;ella.g§erosa,'g5acilaria corticata, Q. edulis andHygnea musciformis and spore release decreased markedly below
20 %o and above 40 %o salinities. The optimum salinity rangeobserved for maximum shedding of spores in the four algae was30-40 %o (Figs. 22-25). Though the optimum range of salinityvaried on monospore production from conchocelis phase in
‘Egrphyra (Yamasaki 3; g;., 1957), monospore release inAcrochaetium endophytium (White and Boney, 1969), formation
and discharge of swarmers in.§;g§ fasciata (Mohan et al., 1972),tetraspores output in Graoilaria corticata, Q, gggggggi,Gracilariopsig sjoestedtii, and Hypnea musciformis (Subba
Rangaiah.g§ g;., 1975; Subba Rangaiah, 1978 and UmamaheswaraRao and Subba Rangaiah, 198O).E1Gelidiumppusillum, Pterocladia
heteroplatos and Qelidiopsis variabilig (Umamaheswara Rao and.Kaliaperumal, 1983), and Dictyota dichotoma (Umamaheswara Rao
and Sanjeeva Reddy, 1982), the general trend observed in theformation and release of reproductive elements in differentsalinities in these algae was similar‘to that observed inthe present study on Gelidiellg acerogg, Gracilaria_g9rticata,‘Q. edulis and Hypgggflmusciformis. The optimum range ofsalinity found in the laboratory experiments is nearer to
the annual range of 28.28 - 36.2§Z§alinity recorded duringthe present (Figs. 55-57) and earlier investigations (Prasad,1954 and Jayaraman, 1954) for the inshore waters of Mandapam
area. These findings indicate that the salinity of the
94
sea water in the nearshore area of Mandapam is suitable forliberation of maximum number of spores in all months of the year.
LIGHT INTENSITX
In some species of the order Gelidiales spore output studieswere carried out by Katada (1955). He observed that theshedding time in the case of tetraspores was earlier in thelight than in the dark, but carpospores were not affected.Rao (1971 a) also reported similar observations in Gelidiellaacerosa, where spore liberation was slightly promoted bylight. Monospores were obtained in the light intensities of19-500 lumens/sq.ft. (205—5382 lux). The sporesbleached anddied soon after release between 480-500 lumens/sq.ft.(5167-5382 lux) and spore output was not found in dark.Massive spore output over long periods was seen in the light_intensity range of 50-110 lumens/sq.ft. (538-1184 lux).In.Monostrgmg (0hno, 1982) liberation of gamete was foundbetween 500 and 5000 lux and at higher intensities of 10,000and 20,000 lux no liberation of reproductive bodies occurred,although gamete formation.was ohserved. However,_gameteliberation was seen at 10,000 and 30,000 lux light intensitiesin.Monostroma ggtidum (Ohno and Nozawa, 1972).
Subba Rangaiah (1978) and Umamaheswara Rao and Subba Rangaiah
(1980) reported maximum shedding of tetraspores in Gracilaria£9PtiCata, Qgggilaria textorii and Gracilariopsis sjoestedtii
95
in complete darkness. The values obtained at 750 3 50 luxwere slightly less than those obtained in darkness and thespore output decreased gradually at 1500 and 2000 lux lightintensities. But in Hypnea musoiformis spore output was higherat 750 t 50 lux light intensity than in darkness and the sporeproduction declined in the other two light intensities.Umamaheswara Rao and Kaliaperumal (1983) found that peak
discharge of_tetraspores in Gelidiug_pusillum, Pterocladigheteroplatos and gglidiopsis_Xariabilis at 500 lux. From1000 lux the spore output decreased with increasing illuminance
in Gelidiumgpusillum and Gelidiopsis variabilis and completeinhibition was seen in these two algae at 5500 lux. InEterocladia heteroplatgg the peak output was at 500 lux.Changes observed between 1000 to 3000 lux were not marked
and lowest values were obtained at 4500 and 5500 lux,light intensities.
In the experiments carried out with the fruiting thalli ofGeligiella acerosa, Gracilaria corticata,.Q._ggg;;§ andHxpnea muspiformis at different light intensities, in thepresent study spore release occurred from 0 to #000 lux.This observation agrees with those of Ohno (1972) on gameteliberation in.Monostroma, White and Boney (1969) on monospore output in Acrochaetium endophytium and Umamgheswara
Rao and Kaliaperumal (1983) on tetraspore shedding inGelidium pusillum, Pterocladia_h§teroplatos and Gelidiopsis
96
gariabilis. gubba Rangaiah (1978) and Umamaheswara Rao and
Subba Rangaiah (1980) observed peak discharge of spores incomplete darkness in Gracilaria gorticata,ԤN'textorii andGracilariopsis sjoestedtii whereas in the present studymaximum liberation of spores was observed at 500 lux in
gglgdiella acerosa, Graoilaria corticata and.§. edulis(Figs. 26-28). The results obtained on these three algae inthe present study confirms with the results of UmamaheswaraRao and Kaliaperumal (1983) for ggligium pgggllgg,.E£2£92le§ie 2££§£9R$§E9§:3nd.§E$£§$9R§i§ variabi}iPis in
which maximum tetraspore output was observed at 500 lux.
Peak output of spores was found at 1000 lux in.gZpggamusgiform;§_gr0wing on.Mandapam coast (Fig.29) and it is
similar to that observed for Hypneg musciforgig growing onVisakhapatnam coast (Subba Rangaiah, 1978 and Umamaheswara
Rao and Subba Rangaiah, 1980) where maximum spore output
was seen at 750 :_50 lux. In.Mono§§roma.Q;§iggg gameteliberation was seen at very high light intensities rangingfrom 10,000 to 30,000 lux (0hn0 and Nozawa, 1972). But ingggggggggg inhibition of spore emission was found at 10,000and 20,000 lux (Ohn0, 1972) and in_§elidium pusillum and§g}g§gpp§i§,g§gggb$l;§ at 5,500 lux (Umamaheswara Rao and
Kaliaperumal, 1983). In.§§g;Qg;ggg§;ggfigggplafigg lowestvalues in spore release were obtained at high lightintensities of 4,500 and 5,500 lux (Umamaheswara Rao and
Kaliaperumal, 1983). Similarly in the present investigation also minimum number of spores released was seen inthe four algae at 4,000 lux light intensity.
222129593322 =
The relation between sporulation of seaweeds and sea watertemperature was studied by Suto (1950 a and b). Accordingto Suto (1950 a) spore shedding in seaweeds takes placewhen the sea-water temperature has reached a level optimum
for each species. The spore liberation in.gg;iggpg;gg§£§g;takes place daily in the afternoon, probably because theright temperature is reached then. He further stated thatshedding season started in_§g;;gigg when the sea watertemperature rose to 20° C for carpospores. There was anoptimum temperature range for shedding of spores andabnormal temperature delayed or hastened shedding by about20 days. The direct evidence of temperature influence onspore shedding was later supported by Katada (1955) alsoworking on Gelidiales for Qgligium amansii, who observedthat the higher the temperature of the preceding night,the earlier the shedding of spores the following day. Theshedding time of tetraspores and carpospores was notrestricted to the afternoon of a given day but varieddepending upon the temperature of the sea water in thefield. It was also found that there was no change in the
shedding time in the next day when the water temperaturewas less than 25° C. In the experiments conducted byFukuhara (1957) with ;ridQphycu§_gornuccpiae, the diurnalperiodicity in the shedding of spores was not affected bywater temperature. Shedding of tetraspores and carposporesin.ggl;g;gm was observed when the sea water temperature roseto 20° C-and 24° C respectively (Suto, 1950 b). Theoptimum range observed for spore liberation in differentspecies of Eggphxra varied from 10-250 C (Kurogi andAkiyama, 1966) and from 1-15° C (Kurogi _&3_;L_., 1967).
Kurogi and Akiyama (1966) examined the effect of watertemperature on the liberation of monospores on 6 species ofP<>r-h ra» namely 2- .’9..e;£.1.<:.r.e»* .2» l;.u2.ie_da.il .2» xe.2.=.9.ez2.$.é.-.§:
§. angusta,_§, suborbiculata and 2; pseudolinegrigaMonospores were liberated between 100 and 25° C in"
E. §ggg§g,;2..x§gggg§;§ and_§. ggggggg. The liberation waslittle or not seen in.§, gggggggg, E. suborbiculata andE; pggggglggggggg at these temperature ranges. Kurogi.33 3;. (1967) noticed differences in the optimum temperatureof monospore liberation between the conchocelis of the
autumn and spring plants of Eggphxgguumgiligglig, althoughexperiments flnbh these plants were carried out at the sametime. Abundant monospores were liberated at 5°, 10° and15° C in monoecious spring plant. There was no significantinfluence on spore liberation in the temperature range of
-CnZ5%q6
99
15~25°mc in Gloiopeltis tenax and 10° c—2o° c”in”c1diqpe1t;§furcggg. At temperatures above or below these rangesliberation was delayed and the number of spores shed alsodecreased (Matsui, 1969).
Tetraspore shedding varied at 5 different temperaturesranging from 15° c to 35° C in Graoi1ar;g_ggrticatg,Gracilaria textorii and Hypnea musciformig (Subba Rangaiah,1978 and Umamaheswara Rao and Subba Rangaiah, 1980). There
was no spore discharge at temperatures below 15° C and above350 C and peak shedding was seen at 30° C. The temperaturebetween 25° C and 30° C was considered as optimum for dischargeof maximum number of spores in these three algae. Thetetraspore output varied at 9 different temperatures rangingfrom O0 C to 45° C tested with the tetrasporic thalli ofGelidigm_pusillum, Pterocladia heteroplatos and Gelidiopsisvariabilis (Umamaheswara Rao and Kaliaperumal, 1983). InGelidium_pusillum spore output increased from 15° C with peakshedding at 25° C. At 30° C and 55° C the spore output
declined rapidly. Inlgggggglgdggiggteroplatgg minimumshedding was obtained at 10 and 15° C and also at #00 Cshowing its ability to liberate reproductive elementsat low and high water temperatures. Between 20 and 35° C
the changes observed inifieteroplatos were similar to thoseof Gelidium_pusillum with peak discharge of spores at 25° C.In contrast, in.§§lidiopsi§ variabilis spore output was
100
seen only at three temperatures (25, 50 and 35° C) with amaximum at 50° C. There was a sudden fall in the quantityof spores liberated by this algae at 35° C. Peak discharge
of spores was.found at 25° C ingpusillum andfheteroplggggand at 30° C in%§ariabili§ showing that the temperaturesbetween 25° C and 30° C was optimum for maximum liberation
:of spores in these three species. In the experiments conduqfified with Dictyota dichotoma at different temperaturesranging from_1O to 45° C, maximum output of tetrasporeswas seen at 30° C.’ There was no shedding of spores fromthe fronds subjected to 10° C, 40° C and 45° C temperatures.When compared to the other temperatures tested, the valuesobtained at 25° C was slightly high indicating that temperatures from 25 to 30° C were favourable for maximum liber
ation of tetraspores in_Diotyqta dichgtoma (Umamaheswara Raoand Sanjeeva Reddy, 1982).
In glgggpgltgg spp. peak output of spores was reported from~10° C - 25° C (Matsui, 1969). The optimum range of
temperature for the species of Qglggggm, Porphxra andgloiopelggg was found to be low as compared with those forthe maximum release of spores in_§gl;g;g;;g acerosa,Gracilaria 9.9.r.:9.i.2_§:c_e. 9- .e.<_1.t.:.;L..J;§. and .P.ixp.n..es masciformis
growing in the vicinity of Mandapam. The optimum temperaturerange for maximum liberation of spores in these four specieswas 250 C - 300 C (Figs. 30~33) as reported for the release
101
of tetraspores in Gelidium pusillum, Pterocladig heterqplatos,Gelidiopsis variabilis, Gragilaria_gorticata, Q, textorii,Hxpnea musciformig and Dictyotg dichotoma growing atVisakhapatnam coast (Subba Rangaiah, 1978; Umamaheswara Rao
and Subba Rangaiah, 1980; Umamaheswara Rao and SanjeevaRaddy, 1982 and Umamaheswara Rao and Kaliaperumal, 1983).
This temperature range coincided with the annual range ofsurface sea water temperature recorded for the inshore
waters of Mandapam during the present investigation (Figs.55-57) as well as in the earlier observations (Prasad, 1954;Jayaraman, 1954 and Umamaheswara Rao, 1972 b).
Photoperiod:
The effect of day length on sporogenesis has recently beenreviewed by Dixon and Richardson (1970). In Bangig fusco:pyrpurea, the Chonchocelis phase produces monosporangiaonly in short day photoperiod (9L : T5 D, 12L : T§'D, or9 L :'9'D). Literature available also suggests that daylength or photoperiod influence the spore production.In.§g£pgx£g.ggggg§ (Iwasaki, 1961) monospore formationand release of fertile monospores were induced by short day
pcondition of 8-11 hours light period. Work on photoperiodinduction and liberation.of monospores from the conchocelisphase Of 5 species of gggpgxgg were carried out by Kurogipand Suto (1962). The adequate dark period for the liberationof monospores in Porphgra tenera, E. kuniedgi, P. yezoensis,
102
3. pseudolinearis and‘£. angu§tg'was 12-16 hours (Shortday condition) while Kurogi and Suto (1967) observedmore liberation of monospores in Porphyra ggbilggglggunder long day conditions. The Acrochaetium endophyticum(White and Boney, 1969) plants kept under long day conditions (18 i66'LD cycle) produced large number of monospores at all light intensities ranging from 30-590lumens/sq.ft. (323-5382 lux). In Monostron3_§1_, Ohno (1972)found long day treatment (18-24 hours light period at3000 lux) resulted in faster gamete liberation than frondsgiven short day treatment at the same light intensity orfronds maintained under natural day-night regime. InGelidiella acerosa (Umamaheswara Rao, 1974 a) spore outputwas found at O + 5?, 8 + T6, 16 + E and 24 + 5'light anddark periods and maximum shedding was observed at 16 + 6 LD
cycle. The present observations made dn.Gelidiell§ acerosaagree with the findings of Umamaheswara Rao (197h a).
In Qracilaria corticata release of tetraspores and carpospores was observed in different light and dark periods
(Umamaheswara Rao) 1976). The tetraspore output was maximumin O + §E LD cycle and with increase in the light periodthe spore shedding gradually deceased. The carpospore outputvaried under different light and dark cycles and maximumnumber of spores was liberated in.4 + 55 LD cycle. Similarlymaximum shedding of tetraspores was also observed by Subba
103
Rangaiah (1978) and Umamaheswara Rao and Subba Rangaiah
(1980) in.Q5§g;;ar;§ corticata, ggggiggria textorii,Gracilariopsis sjoestedtii and Hypnea musciformis atO + E5 LD cycle, and the spore output declined with increasein the duration of photoperiod. The values obtained at4 + 25 LD cycle were also higher than in other photoperiods.
The above findings is in confirmg¥§en with the presentinvestigations made on three species of the order Gigarti_r__1,ales (Gracilsrie corticats: 9.» _a~:.1_s.l.J',+.§ and Exaees .I1.12.s«..<>_i;.£..s>_;?.n.1fi.;_§)
Recent studies made on the combined effects of light intensity
and photoperiod have indicated that the amountfienergy receivedby the plants affects their growth and reproduction andwhile reviewing the earlier work, Dixon and Richardson (1970)described the interacting effect as photosynthetic effect.Murray and Dixon (1973) reported the effect of light intensity and photoperiod on apical cell division in Pleonosporiumsquarrulosum. Rate of cell division was found to be affectedby both light intensity and light periods of 16 + 8'LD and8 + To LD and increase in cell division was found upto avalue of 558 mean daily Illuminance (Illumination in luxmultiplied with photoperiod in hours per day). The combinedeffects of illuminance and duration of light on the shedding
of tetraspores from Gelidium pusillum, Pteroqlgdig hetero?platos and gglidiopsis variabilis was recently studied byUmamaheswara Rao and Kaliaperumal (1983). At 500 lux spore
10fl
output increased gradually and peak shedding was observedunder long day conditions between 1653 and 24 : 5'LDphotoregimes. At higher illuminance maximum sheddingwas seen under short day conditions. Maximum sheddingof spores at 2000 and 3000 lux in Gelidium_pusillum andPterocladia heteroplatos was observed at 8 : 73 LD cycle.At 4500 lux spore output was high in these two speciesunder 4:20 LD cycle. (Although variation exists inrelation to intensity and duration of light, spore output was low at higher illuminance values as observed inthe data collected under continuous light. The variationsin the mean values of spore shedding were observed indarkness (0:25 LD) and are due to the use of materialscollected at different times of the year). Thus therole of light (intensity and photoperiod) in inducingspore shedding is still obscure and that further workneeds to be done to investigate this relationship inother seaweeds of economic importance.
In the combined effect of light intensity and day length(light and dark cycles) the earlier works in gggphyggspp. that indicate the release of reproductive bodies
were induced by short day conditions onlyfby Iwasaki,(1961) and Kurogi and Suto, (1962) in Gracilagia corticataby Umamaheswara Rao, (1976) and Subba Rangaiah, (1978)
in Gracilgria textorii, Gracilariopsis sJoestggt;i.and
105
gypggg mgsciformis by Subba Rangaiah, (1978) and in longday conditions in Porphyra umbilicalis by Kurogi and Suto,
(1967), in Acrochaetium endophytigm by White and Boney,(1969), in Monostroma by Ohno, (1972) find in Gelidiellaacerosa by Umamaheswara Rao, (1974). In _1-I_3_a_ng_i_._a _:§_}_1_§__c_o_~
purpurea, the ghggghggglig phase produce monosporangia(Dixon and Richardson, 1970) only in short day photoregimes (9L : 15D, 12L : 12D, or 9L : 9D). Similarlythe phase of .F_’.<2rp.1:1:z.;".e .i..:.§.I3.9.1;e Produced mo no
sporangia (Kurogi and Suto, 1962) only in a short dayphotoregimes. The same was also observed‘in.g;§g§_g§;;fornia which produces monospore under 8L : 16D photoregimes(Scott and Dixon, 1971).
From the above it is seen that in the first two genera(§,fuscopurpurea_& E. tenera) the response has been shownto be truly photoperiodic since sporogenesis is inhibitedby light interruption in the middle of the dark period.In,gg£gghgg3;gm pggtipgtgg tetrasporogensis occurs onlyin photoperiods of 10 hours or less (in a 24 hour cycle)but exposure to light in the middle of the dark period(West 1968) was not inhibitory, hence in this case theresponse does not appear to be truly photoperiodic. Onthe other hand Pseudoglpiphlggg gggguggg showed (Ramus,1969) a complex interaction of temperature—light intensityand light duration in tetrasporogenesis. In photoperiods
106
of 16L : §E, the Maximum spore production was at a temperature
of 200 C and at light intensity of 2,700 meter candles butwhen the temperature was reduced to 150 C and the photoperiodand light intensity remained the same, spore production wasreduced. When the thalli were grown at 15° C under aphotoperiod of 16L : 50 and at 1,100 meter candles lightintensity, spore production was totally inhibited. Thus daylength is reported to be an important factor in the production
of spores of some marine algae. In his studies on.§grph1§§tenera, Kurogi (1959) observed that daily photoperiods of 8-11hours not only induced monospore production and maturationbut also induced the release of the monospores. In Gelidiumpusillum peak spore output was observed at 16L :'§b in 500 luxand at 8L : 1'57) in 2000 lux light intensity. At 4500 lux lightintensity high spore shedding was seen in complete darkness(OL : Em) than in AL : 20D. Similar trend in spore sheddingwas observed at 500 and 4500 lux light level in Pterocladiaheteroplatos and at 500, 2000 and 3000 lux in_gelidiqp§i§gariabilis (Kaliaperumal, 1978).
In the present study, peak liberation of spores was notedunder long day conditions at low light intensity and at highlight intensity under short day as well as under long daylengths. In gelidiella acerosa maximum tetraspores wereobtained at 500 lux light intensity under a photoperiod of
24L : 00, at a laboratory room temperature of 30° C. -At2000 lux light intensity maximum spore output was obtained
107
at 16L : Eb, 20L : Eb photoperiods and at high lightintensity (4000 lux) maximum spore shedding was seen in shortday length of AL : 56D as well as in complete darkness.Similar results were obtained for tetrasporogenesis andcarposporogenesis in Qracilaria gprticata,.§._gggl;§ andHypnea musciformis. Thus day length appears to be animportant factor in the spore production and spore shedding.Their interacting effects show that the amount of lightenergy received by plants influences spore release and thisperiodicity in the discharge of spores is due to thephotosynthetic effect as has been described by Dixon andRichardson (1970). Similar photosynthetic effect was observedby Murray and Dixon (1973) in the division of apical cells inPleonospgrium squarrulosum. From the present study on sporeshedding it may be presumed that the Mean Daily Illuminationis an important factor in marine algal growth and reproduction.
_gFoRMI NATION? or sPoR§_s_
Information available in the literature indicates that therate of germination of red algal spores varies in different
.;, :theueh aruitiefimeans Of the Year 3339 in Aplants were found throughout the year, spores germinated inthe spring and early summer (Barilotti and Silverthorne, 1972).Another similar observation was reported by Prince andKingsbury (1973) for seasonal variations in the growth.of
germlings of_phondru§ crispus. Data obtained in the present
108
study on Gelidiella acerosa do not agree with the abovefindings as the germination of spores were observed throughoutthe year. There was no particular season for spore germinationas the fruiting plants were present throughout the year ofstudy and spores liberated settled at the bottom of the petridishes and germinated soon after settlement. This wasobserved during the spring tide periods throughout the yearof study. Germination of tetraspores and carpospores ofGracilaria g9rticata,‘Q. edulis and Hypnea_musciformis wasaalso observed throughout the year of study, except in thecarposporic plants of Hypnea musciformig which were availableonly in some months at stations II and III.
In the other gelidiaceous algae e.g. Gelidium pusillum and_Eterocladia heteroplatos highest number of dividing sporeswere observed during and after the peak growth period(Kaliaperumal, 1979), whereas in Qelidiopsig variabilisspores did not germinate within 24 hours. In the presentstudy the spores of Gelidiella acerosa germinated soon afterliberation and settlement at the bottom of the petri dish.The germination rate was higher in Gelidiella than thatobserved in species of the order Gigartinales e.g. Gracilaria_9orticata,.§. edulis and fiypnea musciformis collected fromthe vicinity at Mandapam region. The tetraspores andcarpospores also germinated within 24 hours of liberationin the above three species of the order Gigartinales.'
109
There was no seasonality in germination of spores as thereproductive plants of tetraspores and carpospores werefound throughout the year and plants were collected duringthe spring tide periods. The spores (tetraspores andcarpospores) germinated a few hours after liberation fromthe parent thallus. These observations suggest that duringthe two spring tide periods in each month maximum sporeoutput and germination takes place than at other times.
Seasonal germination of spores was observed by some'workers.Barilotti and Silverthorne (1972) reported seasonal germinationof spore in Gelidium robustum from Baia California, in whichjuvenile thalli were observed on artificial substrata only inFebruary. Though fruiting plants were seen throughout the‘year, the spores released during the spring and early summerwere successful in producing young thalli on new substrata.In Chondrus crispus Prince and Kingsbury (1973) observedlow sporulation and growth rates in winter and springmonths and maximum sporulation and growth rates in earlyfall. Subba Rangaiah (1978) observed monthly variationsin the number of dividing tetraspores and carpospores inGracilaria corticata and Hypneagmusciformis released on thefirst day (within 24 hours) from December 1974 to April 1976..The germination rate varied from a minimum of 11 % to a
maximum of 60 % in the two red algae and highest numberof dividing spores were seen during the peak periods of
110
spore shedding from December to February and from Juneto August.
Some preliminary data were collected on the percentagefrequency of germination of tetraspores and carposporesof Gelidfgg pusillum and tetraspores of Pterocladiaheteroplatos by Kaliaperumal, (1979). He found thatthe germination rate of tetraspores in Gelidium pusillumvaried seasonally from a minimum of 0.3 % to a maximumof 13.6 % and that of carpospores from 0.2 to 39.1 %.Highest rate of germination (within 24 hours period)was observed in tetraspores and carpospores of ggligigmpusillum in February and in general high germination ratewas observed during and after the peak growth season. InPterocladia hetgggplatos the germination rate of tetraspores was less than that observed in Gelidium pusillumand data obtained in different months ranged from 0.5 to4,1 %. The germination rate was less than 1 % exceptin the Ebnth of September. Only 4.7 % of carposporesof Pterocladia heteroplatgg germinated in experimentsconducted during September. In other months there were
no divisions in the carpospores liberated. Dividing sporeswere not observed (within 24 hours) in Gelidiopsisvariabilis during the two and a half years‘ seasonalstudy on spore output (Kaliaperumal, 1979). During thepresent investigation on germination of spores (tetraspores and carpospores of the four selected species of red
111
algae), it is observed that the liberated spores do not~follow any regular seasonal periodicity in the germinationas reported by N. Kaliaperumal (1979) Subba Rangaiah(1978)
Spore germination and growth.have been shown to beinfluenced by environmental factors such as desiccation,salinity, light and temperature. In experiments conductedat room temperature with Gelidium pusillum, maximumgermination rate (18.5 %) was seen in controls and the
germination rate decreased with increase in the durationof exposure upto 45 minutes (Kaliaperumal) 1979).
Germinating spores were found inwgterocladgg heteroplagosbetween 15 and 60 minutes exposure periods and in fronds
exposed for more than one hour spore germination was notseen.(Ka1iaperumal, 1979); In the present study of thespores easily germinated in the fronds(fertile) kept underesubmerged conditions, than those fronds which were exposedto air.
In Gracilaria corticata dividing spores were found in
salinities ranging from 20 to 40 %o with their maximumnumber at 30 %o andfhypnea musciformis though peak outputwas found at 20 %o the rate of spore germination wasmaximum at 30 %3 (Subba Rangaiah, 1978). In different
salinities tested with the tetraspores of'gg;;g;Emgpusillum and Pterocladia heteroplatos germination of spores
112
‘was found only in Gelidium pusillum at 20 and 30 %o and thevalues were 1.0 and 0.7 %o respectively.
The effect of light intensity on the germination ofcarpospore in'§;§g;;g§;a*verrucosa was studied by Jones(1959 a). Development of carpospores was not observed byhim when the carpospores were exposed to high lightintensities for 4 to 5 hours daily. Spore germinationwas also observed in Acrochaetigg endophyticum by Whiteand Boney, (1969) under 19-420 lumens/sq.ft. (205-4521 lux)and it was not found at dark and 480-500 lumens/sq.ft.(5167-5382 lux)Q In Gracilaria corticata and Hypneamusciformis though peak production of spores was observedin dark and low light intensity, germination rate increasedfrom 0 to 1500 lux intensity and thereafter slight decreasewas seen by Subba Rangaiah, (1978). The rate of germinatingspores was high between 1500 and 3000 lux light intensitiesin Gelidium pusillum and the values obtained at 500 and1000 lux light intensities were less than those obtainedin dark. In light intensities above 3000 lux there wereno dividing spores. Similar trend wassnot observed inPterocladia heteroplatos by Kaliaperumal, (1979);
113
Katada (1949 and 1955) studied the effect of temperatureon the germination of carpospores and tetraspores inGelidium amansii. The optimum range of temperature underexperimental conditions was found to be about 24 OC — 26 OC.Below this range of temperature the germlings developedslowly but were healthy in general, while above 28 °c thesporelings died quickly. In lridophxcus cornucopiae theupper and lower limits of temperature for the germinationof carpospores as well as tetraspores were about 3 O - 23°Cand the optimum temperature was 70 - 15°C (Fukuhara, 1968).
The effects of temperature and salinity on the growth ofsporelings of Eelidium amansii were studied by Ohno(1969).He reported that the growth decreased above and belownormal sea water concentration and reduction in growth wasmore at 25 0C than at 10 °C. Spore germination was notseen at low temperatures of 10 OC and 15 0C in Gracilariapggggigggg and Hgpgea muscifgrmis. Dividing spores werefound between 20 OC and 30 °C and rate of germination was
maximum at 30 OCa These temperatures agree with thesurfaoefiaater temperatures recorded along the Visakhapatnam
003st (Subba Rangaiah, 1978). In experiments conductedat different temperatures though the values were irregular,germination rate increased from low to high temperaturesboth in.gglidium_pusi1lum and gterocladia heteroplatos
114
and high values were obtained at 30 OC and 35 OC by
Kaliaperumal, (1979).
In relation to the influence of environmental parameterson the germination and further growth of germlings, in
Gelidium amansii was observed by Ohno, (1969)o He showedthat the growth of sporeling decreased above and below thenormal salinity of sea water. In Gracilaria_verruc9sa,Jones, (1959) did not observe development of carposporeswhen exposed to high light intensity for 4 to 5 hours daily.White and Boney (1969) also did not observe germination ofspore in Acrochaetium endophyticum in dark as well as inhigh light intensities of 480-500 lumens/sq.ft. (5167-5382lux). Similarly Katada, (1949 and 1955) reported anoptimum range of 2# CC - 26 CC for the development ofcarpospores and tetraspores in Gelidium amansiis From thepresent investigations on spore germination given in Figs.45 to 54, it may be pointed out that submerged conditionof the fronds, sea water salinity around 20 to 35 ppt.,light intensity of 500 lux to 3000 lux and temperatureOf 25 CC to 35 0C are favourable for germination of sporesin gelidiella aaerosa, Grgc1;aria corticata,.Q. edulisand_HXpnea musciformis. Complete darkness, short dayconditions as well as long day conditions were also suitablefor spore germination as observed in the present studies.
115
These are in coniirmity with the findings of Katada (19h9and 1955); Jones, (1959); Ohno (1969); White and Boney(1969); Burns and Mathieson, (1972). In Gigartina stellatathe optimum germination of spores occurred at 30 %oS andmaximum growth rate at 25, 30 and 35 @o salinity. Thespores did not germinate at 15 %c, but showed some germination and growth at 50 %o as is found in the presentobservations made_on the germination of selected red algae,is given in (Figs. 47-48), which also agree with theobservations made by Subba Rangaiah (1978) on some speciesof G$gartinales, and by Kaliaperumal, (1979) on two speciesof Gelidiales.
Mode of spore germination 3
The divisions in the germination of spores were observedin.the tetraspores and carpospores of the three members ofthe order Gigartinales studied. Gracilaria and Hxpnea showeda similar pattern of segmentation of spores In the orderGelidiales (Gelidiella acerosa) where a lateral protuberanceis seen where the protoplasm from the original spore migrates
into the protuberance and iurther divisions were observedin the lateral protuberance from which adult thallusdeveloped. Thus type of spore germination found inGelidiella gggggg is characteristic of the order Gelidiales.It is described as 'Gelidium type‘ of germination(Fritch 1945)
116
In Gracilagig corticata,g§. edulis and_§ypng§ImE§g;;g§Q;§(Gigartinales) the protoplasm remains enclosed in theoriginal spore and divides by a wall perpendicular to thesubstratum, giving rise to two cells, and then by theformation of a wall perpendicular to the first one givesrise to four cells, forming a quadrant. The four celledsporeling later by further division forms a multicellularstructure. This later on forms a disc-like body fromwhich rhizoids, unicellular hairs and young thalli formedThis type of division observed in Gigartinales is describedas 'Dumontia type’ by Chemin (193fXtFrmtsch(fl945).Fragmentary studies on germination of red algal spores havebeen shown to be of diagnostic value in determining thetaxonomic and phylogenetic relationships (Katada 1955;Chihara & Kamura, 1963). Similar types of observationswere reported in the order Gigartinales (Rhodophyta) bySubba Rangaiah (1976) in the germination of tetraspores
and oarpospores of Gracilaria corticata and Hypneamusciformis occurring in the Visakhapatnam coast. Studies
by Jones (1959) on Gracilaria verrucosa, Feldmann (1962)on Corydelgladia erecta, Burns & Mathieson.(1972) on
Chondrus.grispus have confirmed the 'Dumontia3 or"mediate discalis“ type of spore germination in the abovealgae.
117
Hgdrography 3
The environmental parameters (e.g. sea water temperature,salinity; dissolved oxygen and pH) vary seasonally andthis (Conoover 1964) cause some species to disappear andreappear according to the season and optimum conditions;
Temperature :
Temperature is one of the most important factors affectingthe growth, reproduction and many other activities in theliving organism. The mean temperature of a given locality,the seasonal and the annual variations do play veryimportant roles in the littoral, or intertidal to sub~tidalzones, varying in seasonal and geographical distribution,pattern of many marine algae. The effect of temperatureon the validity and growth of seaweeds and developingsporelings has been investigated by several workers. InScinaia fercellata and fielminthoclagig purpurea thecritical upper temperature limit is 35 DC to 40 OC.(Chemin 1937). He further reported that the tetrasporesof Gelidium latifolium tolerated a temperature maintainedat 39 °c for 30 minutes; but the carpospores ofCallophyllis laciniata succumbed at 36 0C after 30 minutes.Inlgglggggm amansii the favourable range of temperature forthe development of carpospores is 24 0C to 26 OC and below
118
24 OC development of sporeling is_very slow, whereas above28 °C the sporelings die quickly. In Codium fragile thefavourable temperature for spore germination (80%) (Weber
1968) is 22 CC, whereas as at 8 °C there is only 10% spore
germination. Sporeling from the monospores of Porphyrarequire a temperature ranging from 60 to 14°C for maximumgermination, (Yendo, 1919) while those of tetraspores andcarpospores of.Qhondrus crispus showed optimum growth
at 15°C aid at 2008 there was no germination (Chen andMclachlan 1972). In Gigartina stellata, the growth rateof the sporeling was accelerated with increase in temperaturesto 19 °C (Burns and Mathieson 1972). At relatively hightemperatures the increased respiratory rates may exhaustthe algal food reserves. If this is prolonged the growthof algae would be arrested (Moore 1958). In tropicalintertidal zones, one would expect.that during the period ofRow tide emersion, direct insolation causes algal habitatsto be subjected to relatively high temperatures. Onlyalgae with a high tolerance to elevated temperatures canlive higher up on the shore, hence temperature is one ofthe causal factors of vertical zonation in the distributionof sea-weeds;
119
Figs. 55-57 give the seasonal variations in the monthlyobservations of temperature, salinity, dissolved oxygenand pH of the sea~water in the three collection areasin the vicinity of'Mandapam. These findings suggest thatsea water temperature ranging irm 25.8°C to 31.900,salinity from 28.28 to 36.25%o, dissolved oxygen of 2.67
;to 6.33 ml/l and pH of 8.0 to 8.7 occurring in naturalenvironment are favourable conditions for growth,reproduction, sporulation, spore germination and furtherdevelopment of sporelings to adult thallus of the seaweeds
Generally the ranges of environmental tolerance of marinealgae are_related to the habitats of the algae in whichthey grow. The above findings are in agreement with thesimilar observations made by Ohno (1969) who found that
the growth rates of the early stages of_g;g§ pertusa andPorphyra tenera were maximum at salinities ih theneighbourhood of normal sea water. In Ghondrus crispusthe percentage of carpospore germination and the growthrate of the sporeling varied (Burns & Mathieson 1972)according to the combinedeéfteots of temperature andsalinity.
120
Salinity of the medium in which the marine algae gngalgal sporelings grow is one of the parameters which is
considered to influence many metabolic processes inmarine organisms; A given algal species in questiongrows best in the medium whose salinity is more or lesssimilar to that in their natural habitats.
Ohno (1969) in his observations found that the growth
rates of the early stages of Ulva pertusa and Porphyratenera were maximum at salinities in the neighbourhood ofnormal seawater. The percentage of germinationogarpospore
in Chondrus crispus and the growth rate of the sporelingvaried (Burns and.Mathieson 1972) with the combinedeffects of temperature and salinity, but the maximumpercentage of germination occurred at 11 OC and 35 %osalinity, f-Maximum:-tgyzomth occurred at 19 °C: from 15 %o
to 40 %o, reaching a maximum at 40 %o$, but showed a sharpdrop of zero growth at 50 %o salinity;
In qigartina stellata the optimum germination of thespores occurred (Burns and.Mathieson 1972) at 30 %o Sand 11 EC, whereas.maximum growth rate occurred at 19 OCand 25 %o, 30 and 35 %o salinity. A restricted toleranceto reduced salinities and temperature at 19 50 was observed
in this species by Burns and Mathieson (1972))the spores
123
did not germinate at 15 0/00, but unlike Q. crisgus showedsome germination and growth at 50 0/00. salinity.
Burns and Mathieson (1972) have further reported that someseaweeds could tolerate relatively low salinities underconditions of reduced temperature. For e.g~, the carposporesof Gigartigg stellatg, showed germination and growth at11° C but showed no germination and growth at a temperatureof 19° C.
Salinity has been found to affect rates of photosynthesisand respiration, It has been shown to affect the amountof extracellular carbohydrates diffusing out of some algae'(Foggl1966), In some species of Ulva and Fucus the rateof photosynthesis is doubled when salinity is decreasedby a third (from 35 0/00 to about 10 0/oo salinity),whereas some algae e.g,g§nteromorpha sp.,_§ggg§ vesiculosusand_Egrphyra laciniata, do not show significant changes inrespiration rates with salinity fluctuations, but otheralgae such as.§..serratu§ and ggminaria digjtata showenhanced respiratory rates with decreasing salinity.
In pure cultures of Isochrysis galbana and PrymnesiumIpgggglug_the amount of extracellular polysaccharidesproduced by the algae was increased with reduced salinity.It is a well known fact that the amount of dissolved gasesin seawater is higher under conditions of reduced salinitiesthan at high salinities (Raymont, 1963).
122
Among the dissolved gases, Oxygen is the most importantfor‘hlgal survival. Basically algae are aerobic andphotosynthetic organisms and therefore they require
oxygen for respiration and CO2 for photosynthesis.From the present findings of the dissolved oxygen inseawater, it is seen that dissolved oxygen ranging from2.67 to 6.33 ml/l is favourable for the growth andsporulation of algae, Similarly pH ranging from 8.0 to 8.7are suitable for the growth and sporulation and otherreproductive activities in the seaweeds.
123
CHAPTER - 6
. §_g;vMARY
Studies on sporulation of four commercially important red(sea-weeds)
algae (agarophytes) namely Gelidiella acerosa, Gracilariacorticata,,Qa edulis and Hypnea musciformis growing in thevicinitynof’Mandapam coast were carried out from October 1981to September 1983. During the two years of study; fruitingbehaviour in the natural population of these species was alsoinvestigated. Laboratory experiments were carried out withthe four algaeffigwggfilect information on seasonal aspects ofspore production and diurnal variation of spore shedding.Detailed studies were made under laboratory conditions to know
the effects of some selected environmental factors such asdesiccation, salinity, temperature, light intensity andphotoperiod on spore output in Gelidiella acerosa, Gracilaria.<.e.c.1;.u.lé-.§. and I_i3z;2.es=.e data were alsocollected from the inter-tidal region around.Mandapam area}:The results obtained on all the above aspects are presentedin this thesis.
Population of Gelidiella acerosa, Gracilaria corticata,.g:edulis and Hypnea musciformis occurred throughout the yearALGE EH5 coast oE'Mandapam. Tetrasporic plants of_the fourspecies were observed in all the months of the year} Duringthe period of this study carposporophytes of Gracilaria
124
corticata and.§. edulis were seen continuously throughout theyear, while in Hypnea musciformis they were found only in somemonths from Aug. 1982 to Sept. 1983 at Pudumadam. But at
Kilakkarai the cystocarpic plants were observed almost throughout the year. Cystocarpic plants were not found in thepopulation of Geli§;ella_gcerosa.
Maximum output of tetraspores and carpospores were observed
mostly on the first day of the experiment in the four redalgae studied. The tetraspore output decreased from secondday onwards and rhythmic liberation of carpospores with peakshedding of spores at intervals of different days was alsoobserved. Under laboratory conditions tetraspore output wasobserved for a period of 6-14 days in Gelidiella-acerosa 36-27 days in Gracilaria corticata 3 3130 days in.§. edulisand 3-23 days in Hzpmea musciformis during different months
of the year. Carpospore liberation was found for 6-30 daysin Gracilaria corticata, 10-30 days in_§.‘edg;fls and 2-24days in.Hypnea muscifogmis during the period of this investigation. -Seasonal periodicity was not observed in the liberationof tetraspores and carpospores in the four algae studied andalso marked seasonal changes were not found in the abundanceof stichidia per gram fresh weight of the plant/thalli inGelidiella acerosa occurring at Kilakkarai.
125
Diurnal periodicity in the liberation of tetraaspores with aprominent peak between 2 P.M. and 6 P.M. was observed in
Gelidiella acerosa and there was no change in the time of peakshedding of spores during this study. A definite peak at oneperiod of the day in the shedding of tetraspores and carposporeswas not seen in Gracilaria corticata,.Q. edulis and Hgpneamusciformis and maximum liberation of spores occurred in thesethree species from 10 P.M. to 2 P.M.
In the experiments conducted in the laboratory to study theeffect of environmental factors, spore output declined withincrease in the duration of exposure of fruiting thalli inshade and maximum number of spores were liberated from theplants mostly under submerged condition. The spore outputvaried in different salinities tested, and peak shedding ofspores was seen in 30 %o S in Gracilaria edulis and Hxpneamusciformis, at 40 %o S in Gelidiella acerosa and at 30 %o Sand 40 %o S in Gracilaria corticata. The optimum salinityrange for peak shedding of spores in all the four algae was30-40%oS. Peak sporulation was observed at low light intensity
of 500 lux in Gelidiellgiacerosa,_g;acilaria_corticata and;§.edulis and at 1000 lux in.H2pnea musciformis; Spore outputdecreased in high light intensities. Spore production variedin different temperatures with maximum spore output at 25 0C
in Ge die a acerosa, at 30 °C in Hxpnea musciformis_and at!25 OC and 30 °C in Gracilaria corticata andgga eduliss
126
The germination of tetraspores in Gelidiella acerosa andtetraspores and carpospores in Gracilaria c0rticata,.g. edulisand Hypnea musciformis occurred throughout the year in thelaboratory conditions. But under selected environmentalfactors in the laboratory, maximum number of dividing tetrasporesin Gelidiella acerosa and tetraspores and carpospores inGraoilaria c0rticata,‘g, edulis and Hypnea musciformis occurredin submerged conditions, at salinities ranging from 20-4050,-light intensity from 500 to 3000 lux and temperature 25 to 30°C.
Mode of germination observed in Gelidiella_gggnq§g was that of‘Gelidium type‘ and that observed in Gracilaria corticataeyg.edulis andlHypnea musciformis was that of 'Dumontig type’.
In the sea water samples collected at bi-monthly intervalsfrom October 1981 to June 1983 from the interfidal gcgion atthree localities in the vicinity of Mandapam, temperatureranged from 25.8 to 30.7 °c,,sg;;gg;yy£rpnm28.28%o to 36.25%o,pH from 8.0 to 8.7 and dissolved oxygen from 2.97 to 6.18 m1/litre. The atmospheric temperature recorded in the areavaried from 26.0 0C to 31.5 0C;
127
The present study on sporulation in selected/red algae ofeconomic importance, suggests that the submerged conditionof the fronds, salinity from 25-35%o ; temperature from25-30 °c ; low to medium light intensity (500 to 3000 lux),complete darkness, short day as well as long day conditionsare favourable for spore production, spore discharge, sporegermination, development of sporelings to adult thalli in thespecies of the orders, Gelidiales and Gigartinales, occurringin the vicinity of the coasts of Mandapam.
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CHAPTER - 7
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