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Iodine Content of Some Marine Algae 1

MADELON R. GRIMM 2

THE LITERATURE ON IODINE and iodide con- organisms should both accumulate iodine and
tent of marine algae has been reviewed by lend themselves to modified microbiological
Fritsch (1945), Chapm an (1950), and Smith procedures, numerous species of marine algae
(1951) . To their extensive bibliographies, the were examined . .
contributions of Turrentine (1912) and Cam- The samples studied were collected alive
eron (1914, 1915) may be added. in th e field and air dried in the laboratories
Since man y of the early investigators were of th e collecto rs (see footnotes of Table 1).
interested primarily in the econ omic aspects Upon receipt, these collections were cleaned
of the algae, the y generally disregarded the and redried at room temperature. A weighed
suitability of the species studied for labor- amount (usually 1 gm.) was incinerated to a
atory use. For example, th eir interests were black ash in a porcelain crucible over a Bunsen
concerned with the extraction and ut ilization burner. The .ash was pulverized and mixed
of iod ine for dietar y and medicinal purposes. with 5 milliliters of distilled water and the
The use of this sub starice in fertilizer re- soluble portion separated by centrifugation .
sources was also promoted. The supernatant was shaken up with 1 milli-
In recent years the use of iodine in tracer . liter chlorofo rm, 2 milliliters hydro gen per-
studies with its many applications necessi- oxide, and 1 milliliter glacial acetic acid.
tates an expanded spectrum of available After several minutes, the presence of iodine
species 'of algae to determine which organ- was manifested by a pink to pur ple color in
isms lend themselves to further laboratory the chloroform layer, the intensity of the
study in this field. As far as can be determined, color increasing with iodine concentration.
very little work has been done to ascertain When the nonchloroform layer retained a
the manner in which algae are able to ac- bro wnish tinge, more hydrogen peroxide and
cumulate iodine and retain it against enor- acetic acid were added until decolorization
mous concentration gradients. In search of occurred, and the mixture was reshaken. The
suitable experimental organisms for such stud - color of the chloroform layer was compared
ies and unde r the assump tion that such to color stand ards containin g kno wn conc en-
trati ons of iodine in chloroform . This method
1 Based in part up on data incl uded in a th esis
submitted to Cornell Uni versity in partial ful fillment
is assumed to be a satisfactory means of
of th e requir ements for the degree of M aster of Science. estim atin g the iod ide content of the algal
Th is investigatio n was performe d un der the supe r- ash when iodide is present in concentrations
vision of Dr . .M axwell S. D ory and D r. Walter C.
Muenscher and supported in part by grants from
of over 30 parts per million.
Corn ell U niversity and Graduate Rese arch Fun ds of Quant itative estimations for iodine ob -
Northwestern Un iversity. M an uscript received Febru- tained in thi s investigation are presented in
ary 20, 1952. .
Tab le 1 and are expressed in milligrams per
2 Re search D epartment, Di vision of M icrobio logy,
Ciba Pharmaceut ical Products, In c., Summit, Ne w gram of dry weight of alga and in parts per
Jersey. million . Unused portions of the collections

[318 }
Iodine Content of Algae - GRIMM 319
TABLE 1
IODINE CONTENT OF VARIOUS SPECIES OF ALGAE
MG. 12/ PARTS PER
SPECIES TESTED COLLECTION* SOURCEt GM. ALGA MILLION
Chlorophyta
A nadyomene stellate (W ulf.) C. Ag . . . . . . . . .. . ... B 49-2134 B 0.156 160
Avrainvillea longicaulis (K uetz.) M urr ay & Boodle . B .... B 0.024 20
Bryopsis plumosa (Hu dson) C. Ag . . .. . . 0 0 • 0 • 0 • •• G .... M 0.195 200
. Chaetomorpha linum (M ueller) Kuetz. 0 0 0 0 0 0 0 ••• 0 G .. . . M 0 0
B 49-2203 B 0.110 100
Codium tntertextum Collins & H ervey 0 0 0 ••• 0 •• 0 • 0 B 49- 2139 B 0.300 300
Codium setcbellii G ardne r . . ... . 0 •• • • • 0 • 0 • •• 0 • • 0 H . .. . 0 0.078 80
Dictyospbaeria cavernosa (F¢rsk.) Boergesen . . . . . D 8118 H 0 0
..
0 •

D 8584 H 0.039 40
Halimeda monile (Ellis & Solander) Lamour. ... 0 • • B 49- 2103 B 0.02 20
V Iva expansa (Setch.) Serch. & G ard . . . . . . . ..... H 1054 0 0.059 60
Phaeophyta
A garum cribrosum (M ertens) Bory . . 0 • • 0 • • • • •• • • G 249 M 0.078 80
Alaria marginata Po st. & R upr.. . 0 0 • • 0 0 • 0 0 0 0 • 0 0 H '" . 0 0.059 60
Ascophyllum nodosum (1.) Le ] olis . .... . . . .. . . . .. G .. .. M 0.156 160
Asperococcus ecbinatus (M ert ens) Gre v. . . 0 0 0 0 • 0 . 0 0 G 258 M 0.056 60
Chorda filum (1.) Lamour. . .. .. . . . 0 • • • • • • • • • • • • G ... . M 0.227 230
Chordaria j/agellijormis (M ueller) C. Ag . 0 •••• • 0 • 0 G 283 M 0.313 310
Cladostepbns verticil/atus (Light.) C. Ag . . 0 • 0 • •• • • • G 277 M 1.094 1090
Coilodesme bul/igera Stroemfelt. . . . . . . . . . . . 0 ••••• H 1019 0 0.234 230
Coilodesme californica (R upr.) Kjellm. 0 ••• 0 • • 0 • • • • H 1074, 1075 0 0.25 250
Cystoseira osmundacea (M enzies) C. Ag o 0 • 0 • • 0 0 • 0 • H .... 0 0.117 120
Desmarestia aculeata (1.) Lam ou r. 0 • • 0 • • 0 0 • 0 • • • • • G 287 M 2.5 2500
Desmaresti« munda Serch. & Gard . 0 • • • • • 0 •• • • • • • H 7689 0 0.469 470
Dictyopteris plagiogramma (Mont.) Vickers .... 0 •• B 49-2081 B 0.234 230
D 856 1 H 0.02 20
Dictyosipbon foeniculaceus (H udso n) Grev. : ... . . .. G . . o. M 0.284 280
Dictyota dichotoma (H udso n) Lamou r. . . .. .. . .. . . B 49-2206 B 0.013 10
Ectocarpus siliculosas (D illwyn) Lyngbye . . 0 0 • • 0 •• • G . . o. M 0.087 90
Elachistea jucicola (Velley) Aresch.... 0 • • • • • • • ••• G .. .. M 0.195 200
Fucus evanescens C. Ag ... . .. . . . . 0 • 0 0 • 0 • 0 0 0 0 • ••• G 278 M 0 0
H 1088 0 0.078 80
Fucus evanescens f. oregonensis G ard . . . . 0 0 0 0 • • 0 0 0 0 • H 1077 0 0.078 80
Facas furcat«: C. Ag . . . . . 0 0 0 •• 0 • • • • • 0 •• • • • •• • • H 1078 0 0.117 120
Fucus spiralis 1. . . ... . ...... ... . ... ... 0 ••• 0 0 0 0 G .. . , M 0.120 120
Fucas uesiculosus 1. . . . . .. . ... . .. .. . . . . . . .. . . . . G . . o. M 0.052 50
Haplogloia andersonii (Farlow) Levring 0 0 ••• 0 • • 0 • • H 1064 0 1.25 1250
Hedophyl/um sessile (c. Ag .) Setch.. . . . . . . . . . . . . . H .... 0 1.25 1250
. L aminaria agardhii K jellm .. . . . . . . ... . . . 0 • 0 • • 0 • G 268 M
Y oung th allus .. . 0 0 0 • • • 0 ••• • •••• 0 0 0 0 • • • 0 0 0 • 3.125 3130
Blade . 0 • • •• 0 0 .0 ... . . . . . . .. .. . .
0 •• 0 . 00 •• 0 . 2.083 2080
Stipe ... . 0 • • • • •• •• • ••• • • • • 0 0 • 0 • • 0 0 0 0 .0 0. 00 1.563 1560
H apt era and rhizoids . 0 , • 0 • • 0 0 ••• • • • •• • • • • • • 5.0 5000
L aminaria andersonii Eato n in Hervey . 0 0 •• 0 •• • • 0 H . . .. 0
Blades . . . . . .. .. . 0 ••• o. . .• • ••• 0 0 0 0 0 0 0 0 0 0 • • 0.938 940
Up per stipe .. .. 0 • 0 • • • o. • • • • • • • • • •• • • 0 • •• 0 • 5.0 5000
L aminaria digitata (1.) Edm . . . . . . . 0 •• 0 0 • 0 •• 0 •• G 279 M 1.041 1040
L aminaria saccharina (1.) Lamour.. . .. . . . . . .. . . . H 1065 0
Lower blades 0 • ••• ••• • •• 0 0 • • 0 •• 0 • • 0 0 0 0 •• • •• 1.25 1250
L aminaria sinclairii (Aresch.) An derson 0 • • • • 0 • 0 • H 1058 0
Blade s . . . 0 . 0 0 ••• • 0 . 0 .0 .0 0 •• 0 0 •• 0 • 0 0 • 0 • • 0 • 5.0 5000
H apte ra and rhizoids . . . 0 0 • 0 0 0 0 0 •• 0 0 0 0 • • 0 .0 • 1.875 1880
Stipe . . . . . . . . ... . . . . . . . . . . 0 • •• • • 0 0 • ••• 0 • • • 3.75 3750
Lessoniopsis littoralis (T ilden) R ein ke . . 0 •• ••• 0 •• • • H .. .. 0 0.117 120
M acrocystis integrifolia Bory . . . . . . . . . . . . . . . ... . . H .... 0
Blades . . . . . . , ... .. 0 •• 0 • • 0 • • 0 • • • • • • 0 • 0 • •• • 0.938 940
Axis of branch . . . . . 0 • • 0 • • • • • • • •• • • 0 •• • • • 0 • 0.117 120
320 PACIFIC SCIENCE, Vol. VI, October, 1952

T ABLE 1-continued
MG . 12/ PARTS PER
SPECIES TE STED COLLECTION * SOURCEt GM . ALGA MILLION

Meso gloia divaricata (c. Ag.) Kuetz . G . M 0.240 240


Nereocystis luetkeana (Mertens) Post. & R upr . H . o
Air bladder . 0.938 940
Sterile blades . 0.625 630
Peluetiopsis limitata (Setch. ) Gard . H ... . o 0.156 160
Postelsia palmaeformis R upr. . H o
Blades . 1.25 1250
Stipe . 0.469 470
Ho ldfast. . 1.25 12 50
Pterygopbora californica Rupr . H . o 0.029 30
Ralfsia clavata (Carm .) Croua n . G . M 0.087 90
Sargassum sp . D 8572 H 0.0 2 20
Sargassum filipendula C. Ag . . . . . . . . . . . . G .... M 0.156 160
Sargassum kjellmanianum Ye ndo . H 1067 o 0.02 20
Sargassum natans (1.) ] . M eyen . B 49-2067 B 0.059 60
Sargassum obtusifolium ] . Ag . D 8114 H 0.029 30
Scytosipbon bullosus Saunders . H 1238 o 0.020 20
Soranthera ulvoidea Pos t. & Rupr. . H 1074 o 0.234 230
Sphacelaria cirrhosa (Ro th) C. Ag. . . . . G .... M 0.314 310
Turbinaria tricostata Barton . B 49 -2231 B 0.029 30
Rh od ophyta
Ahnfeltia concinna J. Ag . H 1233 o 0.0 2 20
Ahn/eltia plicata (Hudson) Fries. . . . .. . . H 1219 o 0.117 120
Amphiroa/ragilissima (1.) Lamour. . B ... . B
Coarse type . 0.029 30
Fine type . 0.156 160
Antithamnion uncinatum G ard . H 1145 o 0.02 20
Asparagopsis hami/era (Har iot) Okamura . G 286 M 0.938 940
Asparagopsis sanfordiana H arvey . D 8663 H 1.25 1250
Laurencia spectabilis Post. & Ru pr. . . . . . . . . . . H 1107 o 0.Q78 80
Lophocladia trichoclados (Mer tens) Schmitz . B 50-426 B 0.117 120
B 49-2086 B 0.117 120
A sparagopsis taxi/ormis (Deli le) Colli ns & H ervey . B 50-5 16 B 1.875 1875
Bostrychia scorpioides var. montagnei (Harvey) Post. . B 50- 382 B 0.01 10
Botryglossum /arlowianum (J. Ag.) D e Toni . H .. . . o 0.313 310
Callithamnion baileyi H arvey , . G 203 M 0.234 230
Callithamnion pikeanum H arvey . H .... o 0.039 40
Centroceras clavulatum (c. Ag.) M ont. . B 49 -2 151 B 0.234 230
Ceramium eatonianum (Farlow) De Ton i . H . o 0.039 40
Corallina officinalis 1. . G . M 0.02 20
Cryptopleura brevis G ard . H 7680 o 0.029 30
Cryptosipboni« woodii J. Ag . H 768 5 o 0.117 120
Cystoclonium purpureum (H udson) Batt ers . G 204, 270 M trace
Cystoclonium perpureum var. cirrhosum Ha rvey . G . . .. M 0.039 40
Galaxaura obtusata (Ellis & Solander) Larnour . B 49 -2 144 B 0.117 120
Galaxaura squalida Kje llm . B 49-2070 B 0.059 60
Gelidium arborescens Gard . H 1030 o 0.039 40
Gelidium corneum (H udson) Lamour. . D 8709 H 0.029 30
Gelidium coulteri H arvey . H 1031, 768 7 o 0.117 120
Gelidium crinale (Turner) Larnour . G .. .. M 0.11 7 120
Laurencia obtusa (Hudson) Lamour .
Gree n phas e . B 49-2195 B 0.039 40
Yellow phase . B 49-2194 B 0.1 17 120
M icrocladia borealis R upr. . H 1130 o 0.02 20
Odonthalia fioccosa (Esper) Falken . H 1133 o 0.Q78 80
Odonthalia washingtoniensis K ylin . H 1114 o 0.407 420
Pbycodrys rubens (Hu dso n) Batters . G 285 M 0.313 310
Pbyllopbor« membranifolia( Good. & Woodw.) J. Ag. G .. .. M trace trace
Plocamium pacificum K ylin . H 1191 o 0.029 30
Iodine Content of Algae - GRIMM 321

TABLE I-continued
MG . I.! PARTS PER
SPECIES T EST ED COLLECTION * SOURCEt GM . ALGA MILLION

Plumaria filicina (Farlow) Kuntz .. ...... . . . . ... H 768 1 0 1.25 1250


Plumaria bypnoides (H arvey) K untz . .. . . .. .. . . . . H 1236 0 0.078 80
Plumaria sericea (H arvey) Ru pr , . ........ .... . .. G . . o. M 0.32 320
Polyides rotundus (G meIin) Grev . . . . . . . . . . . . ... . G . . o. M 0.039 40
Polyneura latissima (Ha rvey) K ylin . . . . . . ... . . .. . H 1147 0 0.039 40
Polysipbonia elongata (H uds on) H arvey . ... . . . . . . G • . o. M trace trace
Polysipbonia lanosa (1.) T andy ... . . . . . .. .. . . . . . . G . . o. M 0.438 440
Porpbyra naiadum Anderson .. . . . . . .. . . .. . .. . . . H 1023 0 0.313 310
Pterocbondria woodii (H arvey) H ollenberg .. .. . ... H 1148 . 0 0.078 80
Pterosiphonia biplnnata val. robnsta (G ard.) Dory .. H 1213 0 0.078 80
Rhodomela larix (T urner) C. Ag. .. . . . . . . . . . . . . . . H 1033 0 0. 156 160
Rhodymenia californica Ky lin . . . . . . . .. . . . .. . . . . . H 1196 0 0.078 80
Spermotbamnion turneri (M ertens) Aresch . . . .. . . . . G .... M 0.313 310
Z anardinul« andersonii (J. Ag .) Papenfuss .. . . . . . H 7674 0 0.039 40
Z unardinula filiformis (Ky lin) Papenfuss . .. . . . . . . H 7671 0 0.078 80
Z anardinul« lanceolata (Harvey) De Toni . ... . .. . H 7672 0 0-02 20
H 7673 0 0.059 60
Z anardlnul« Iyallii (Harvey) D e T oni . . . . . . . .. . . H 7675 0 0.117 120
H 1050 0 0 0
* Collector: B, A. J. Bernarowicz; D , M . S. D ory; G, M . R . Grimm ; H , 1. H orow itz .
t Location : B, Berm uda; H, H awaii; M, Woods H ol e, Mass achusetts; 0, Cape Arago, O reg on.

rreated remain for the most part in the her b- and Black (1948). Fritsch (1945) reported
aria of the various collectors. The collector tha t the greatest accumulation of iodine was
and collector's nu mber, where given, are in the actively growing cells at th e base of the
shown in Tab le l. blade. Tab le 1 shows that th e blades of L.
As is shown in Table 1, very little iodine sinclairii contained 5,000, hap tera andrhizoids
was detected in the Chlorophyta, a findi ng 1,880, and stipe 3,750 parts per million. Iodine
which is in agreement with the data of other determinations of o ther species of Laminaria
investigators. Phaeophyta such as Laminaria pro vided simi lar results. Thus, in Laminaria,
and Desmarestia had the highest concentra- th ere appears to be an analogy between the
tions of iodine. H owever, these ge nera are activi ty of gtOwth-reg ulating substances ob -
large forms difficult to m aint ain in the labor- served by Williams (1949) and th e distri bu-
atory. Asparagopsis and Plumaria, bo th R hodo- tion and amounts of iodine prese nt.
phy ta, sto re iodine in cons iderable amo unts Table 1 also indica tes th at two phases of
and have the adva ntage of convenient cultura l Laurencia obtusa contained different amo unts
size. of iodi ne -the yellow ph ase three times as
A variation of io dine conten t was fou nd in much iodine as th e gree n phase. The sam e
different thalli collected from th e same loca l- appears to be true of two gtOwth forms of
ity. For example, one sample of Laminaria Amphiroa fr agilissima. A fine form contained
agardhii collected from Woods Hole showed m ore th an five times as much iod ine as a
no iodine while other specimens of the same coarse form. A potential taxonomic use of the
species from the same vicinity showed an iodide test is suggested here as a suppleme nt
accumulation of 5,000 parts per million. to th e vague morpho logical criteria currently
Turrenti ne (1912) records similar variations used in separa ting the infrageneric taxa of
with L. saccharina. such genera.
In a consideration of the iodine conte nt of The data presented indicate that Asparagop-
the same thallus, marked differences were sis and Plumaria wo uld be the mo st suitable
noted by T urrentine (1912), Camero n (1915), spe cies for further investigation. Th eir adap-
322 PACIFIC SCIENCE, Vol. VI, October, 1952

tation in this study allows the conjecture Vaucheria tburetii W oronin 0


that they may be readily adaptable to methods Phaeophyta
of culture, iodine uptake, plant metabolism, A egira virescens (Carm .) Setch. & Gard . . .M
and tracer studies. Colpomeni« sinuosa (R oth) D erbes & Soc
A listing of algae which ga ve negative lier B
tests far iodine is presented here. These spe - Costaria mertensii J. Ag 0
cies represent approximately 50 per cent of Dictyopteris delicatula Lam our B
all organisms tested . The source of the sam - Dicty opteris j ustii Lamour B
ples is as indicated in Table 1. Dicty ota dentata Larnour B
Dictyota divaricata Lamour B
LIST OF SPECIES YIELDING NEGATIVE Dilophus guineensis (K uetz.) J. Ag , . .B
IODINE CONTEN T Ectocarpus indicus Sonder. H
Cyanophyta Egregia menziesii (T urn er) Aresch 0
Calothrixconfervicola (Roth) Ag . ex. Born. Heterochordaria abietina (Rupr.) Setch . &
& Flah M Gard 0
Lyngbya majuscula "(Dillwyn) Harvey ex. Hydroclathrus clathratus (Bory) Howe B
Gomer' H Ilea fascia (Mueller) Fries 0
Chlorophyta M
Boodlea composita (Harvey) Brand H Leathesia dilformis (1.) Aresch : M
Caulerpa crassifolia (c. Ag .) J. Ag B o
Caulerpa peltata (Turner) Lamour B Padina commersonii Bory H
Caulerpa racemosa var. laetevirens (M o nt .) Padina pavonia (1.) G aillon B
Weber-van Bosse B Padina senctae-crucis B¢rg B
Caulerpa sertularioides (G m elin) H owe B Pocockiella variegata (Lamo ur.) Papenfuss . B
Caulerpa verticillata J. Ag B Punctaria latifolia Grev M
Caulerpa webbiana Mont H Scytosiphon lomentaria (Lyng bye) J. Ag 0
. Chaetomorpha antennina (Bory) Kuetz H M
Cladophora sp H Spatoglossum schroederi (M ertens) J. Ag B
Cladophoropsis membranacea (c. Ag. ) Zonaria zonalis (Lamo ur.) Howe B
B¢r g B Rhodophyta
Codium decorticatum (W oo dw.) Howe B Acanthophora spicifera (Vahl) B¢r g B
Codium fragile (Suringar) Hariot 0 Agardhiella tenera (J. Ag. ) Schmitz M
Cympolia barbata (1.) Lamour B Bangia oermicularis Harvey 0
Enteromorpha clathrata (Ro th) Grev 0 Calloph.yllis crenulata Setch 0
Enteromorpha intestinalis (1.) Link 0 Callophyllis megalocarpa Setch . & Gard 0
Enteromorpha linza var. flexicaulis D oty . . 0 Ceramium paciftcum (Collins) Kylin 0
Halimeda discoidea D ecaisne H Ceramium rubrum (H udson) C. Ag 0
Monostroma grevillei (Thuret) Wittr 0 Ceramium tenuissimum (Lyngbye) J. Ag H
1
Spongomorpha eoalita (Rupr.)' Collins 0 Champia parvttla (c. Ag. ) H arvey M i
Udoteaflabellum (Ellis & Solander) Howe .B Chrondrus crispus (1.) Stackh M
Ulva angust« Setch . & Gard 0 Chrondia sedifolia Harvey M
Ulva f asciata D elile H Constantinea simplex Setch 0
B Cumagloia andersonii (Farlow) Serch . &
Ulva lactuca 1. M Gard 0
H Delesseria deeipiens J. Ag 0
Ulva lobata (K uetz.) Setch , & Ga rd 0 Dilsea californica (J. Ag. ) Schmitz 0
Urospora penicilliformis (R oth) Aresch 0 Dumontia incrassata (M ueller) Lamo ur. . .M
Iodine Content of Algae - GRIMM 323

Endocladia muricata (H arvey) J. Ag 0 Spyridia filamentosa (W ulf.) Harvey . . . . . M


Erythrophyllttm delesserioides J. Ag 0 B
Eucbeuma isiforme (c. Ag. ) J. Ag B Tburetia borneti Vickers B
Farlowia mollis (H arvey & Bailey) Farlow Trichogloea subnuda H owe H
& Setch 0 IJ7ttrdemannia miniata (D rap .) Feldm an &
Galaxattra cylindrica (Ellis & Solander) H amel. B
Lamour. B
R EFER ENCES
Galaxaar« marginata (Ellis & Se lander)
Lamour B BLACK, W . A. P . 1948. The seaso nal variati on
Halymenia f ormosa Kutz H in chemical const itutio n of some of th e
Kylinia sp '. H sub-littoral seaweeds common to Scotland .
L aurentia palisada Yam ada H Soc. Cbem. Lndus., J ottr. 67 : 165- 176.
Liagora f arinosa Lam our. B CAMERON, A. T. 1914. Co nt ributions to the .
L iagora maxima Butters H biochemistr y of iodine . I. The distribution
Liagora valida Harvey B of iodine in pla nt and animal tissues . [our.
H BioI. Chem. 18 : 335-380.
L omentaria baileyana (H arvey) Farlow . .. M - - - 1915. Co ntributio ns to th e biochemis-
Nemalion mttltifidllm (Weber & M ohr) J. try of iodine. II. The distribut io n of iodine
in plant and anima l tissues . [our. BioI.
Ag M
Opttntiella californica (Farlow) K ylin 0
Chem. 23: 1- 39.
CHAPMAN, V. J. 1950 . Seaweeds and their uses.
Petrocelis fr anciscana Setch. & G ard 0
xiv +287 pp ., 20 pls. M ethuen & Co. , Ltd .,
Phy/lophora brodaei (T urner) J. Ag M
Londo n.
Polysiphonia collinsii H ollenberg 0
FRITSCH, F. E. 1945 . The structure and repro- .
Polysiphonia fibrillosa G rev M
duction of the algae. Vol. 2. xiv+ 939 pp.,
Polysiphonia ·harveyi Bailey M 2 map s. University Press, Cambridge .
Polysiphonia paniculata M ont 0 SMITH, G . M ., ED. 1951. M anttal of phycology.
Polysiphonia variegata (c. Ag .) Z anardini .M 375 pp ., 48 pls, Chro nica Botanica Co .,
Porphyra nerocystis Anderson 0 Waltham , M assachusetts.
Porphyra perforata J. Ag 0 T URRENTINE , J. W . 1912. The com po sition
Porphyra umbilicalis (1.) J. Ag M of kelps . Appen dix P . In Fertili zer resources
Porphyra variegata (K jellm .) H us " . .0 of th e United State s. F. K . Camero n . United
Pterosiphonia bipinnata (Post . & Rupr.) States 62nd Congress, 2nd Session, Senate Doc.
Falken 0 190: 217-231, G overnment Printing Office,
Rhodymenia pacifica K ylin 0 Wa shington .
Rhodymenia palmata (1.) Grev M W ILLIAMS, 1. G . 1949. Growth-regulating
Schizymenia pacifica K ylin 0 . subs tances in Laminaria agardhii. Science
.Spyridia spinella Sander H 110: 169.

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