Synopsis of Biological Data on Laminaria Hyperborea

Home , Hyperborea, Laminaria, Laminariaceae, Laminaria digitata, Laminaria hyperborea

FAO Fisheries Synopsis No. 87 FIRM/S87 (Distribution restricted) SAST - Laminaria hyperborea - 7,77(02)002.07

SYNOPI LT OF BIOLOGICAL DATA ON Laminaria hyperborea

Prepared by

Joanna M. Kam

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS e9 ROME, 1971 DOCUMENTS OF THE FISHERY DOCUMENTS DE LA DIVISION DES DOCUMENTOS DE LA DIRECCION RESOURCESDIVISION OF FAO RESSOURCES HALIEUTIQUES DU DE RECURSOS PESQUEROS DEL DEPARTMENT OF FISHERIES DEPARTEMENT DES PECHES DE DEPARTAMENTO DE PESCA DE LA LA FAO FAO

Documentswhicharenotofficial Des documents qui ne figurent pas EstaDirecciónpublica varias series FAO publications are issued in several parmi les publications officielles de la de documentos que no pueden conside- series.They are given a restricted distri- FAO sont publiés dans diverses séries. Ils rarse como publicaciones oficiales de la bution and this fact should be indicated font seulement l'objet d'une distribution FAO. Todos ellostienendistribución ifthey are cited.Most of them are restreinte, aussi convient-il de le préciser limitada, circunstancia que debe indicarse prepared as working papers for meetings, lorsque ces documents sont cités. Il s'agit en el caso de ser citados. La mayoría de or are summaries of information for use le plus souvent de documents de travail los títulos que figuran en dichas series of Member Governments, organizations, préparés pour des réunions, ou de ré- son documentos de trabajo preparados and specialists concerned. These, series sumés d'informationàl'intentiondes para reuniones o resúmenes de informa- are the following: gouvernements des pays membres, ainsi ción destinados a los estados miembros, que des organisations et spécialistes inté- organizacionesy especialistasinte- ressés. Ces séries sont lessuivantes: resados, Estas series son las siguientes

FAO Fisheries Report FIR ¡R (No.) FAO Fisheries Circular FIR ¡C (No.) FAO Fisheries Synopsis FIR ¡S (No.)

Special groups of synopses are iden- Des catégories spéciales de synopses Grupos especiales de sinopsis se dis- tifiedby symbols followed byclassi- sont identifiéesàl'aide de symboles tinguen con las siglas siguientes, seguidas fication numbers based on indexed code suivis des chiffres de classification basés por números de clasificación que se basan of "ASFA ": sur le code d'indexation de la «ASFA»: en las claves de los índices de la «ASFA»:

SAST Data concerning certain species SAST Données sur certaines espèces et SAST Datos relativos a ciertas especies and fish stocks. populations de poissons. y poblaciones. MAST Information on methods and sub- MAST Renseignements surdes méthodes MAST Sinopsis sobre métodos y mate- jects. et des sujets. rias. OT Oceanographic data. OT Données océanographiques, 01 Sinopsis sobre oceanografía. IT Limnological data. IT Données limnologiques. IT Sinopsis sobre limnología. and et y CART Information concerning fisheries CART Renseignements sur les pêcheries CART Información sobre los recursos and resources of certain countries et les ressources de certains pays acuáticos vivos de algunos países and regions (FID/S). et régions (FID/S), y regiones (FID/S).

FAO Fisheries Technical Paper FIRT/T (No.)

Special groups of TechnicalPapers Des catégories spéciales de documents Grupos especiales de documentos téc- are identified by: techniques sont identifiées à l'aide des nicos se identifican por las siglas siguien- symboles suivants: tes:

RE Indexed lists of experts and insti- RE Listes indexées d'experts et insti- RE Listas índices de expertos y de tutionsdrawnfromRegisters tutions tirées des registres tenus instituciones tomadas de los re- maintained by the Fishery Re- à jour par la Division des ressour- gistros que se llevan en la Direc- sources Division. ces halieutiques. ción de Recursos Pesqueros. CB Lists of periodicals, special sec- CB Listes de périodiques, des sections CB Listas de periódicos, secciones es- tions of " Aquatic Sciences and spéciales de la « Aquatic Sciences peciales de la «Aquatic Sciences Fisheries Abstracts (ASFA), and Fisheries Abstracts (ASFA) », and Fisheries Abstracts (ASFA) », specialbibliographies andpa- des bibliographies particulières et bibliografías especiales y trabajos pers concerning documentation des articles sur les problèmes de relativos alosproblemasde problems. documentation. documentación.

MFS Provisionaleditionsof" FAO MFS Editions provisoires des e Manuels MFS Edicionesprovisionales delos ManualsinFisheries Science." FAO de science halieutique s. «Manuales de la FAO de Ciencias Pesqueras ».

Some documents also have another Certainsdocumentsportentparfois Algunos documentos tienen también identification, if, for example, they have d'autresnumérosd'identification,par otra identificaciónsi, por ejemplo, son been contributed to a meeting for which exemple s'ils ont été préparés pour une contribucionesa unareunióncuyos papers have been numbered according réunion dontles documents ontété documentos hansidomarcados con to another system. marqués àl'aide d'un autre système, arreglo a otros sistemas. FAO FiBherieB Sy'eopEi8 8o, 87 FIRM/s87 Dietribution reiotrictod SAST - Laminaria hyperborea .-177(O2)OO2.O7

SYNOPSIS OP BIOLOGICAL DATA QN Laminaria hypex'boroa

Prepared by

JOANNA M. KAIN (Mra. N.S, JONES) Marine Biological Station Port Erin, Isle oMan U. K.

FOOD All]) AGRICUJJPURE ORGANIZATION OF THE UNITED NATIONS Rome, December 1971 PREPARATION OF THIS SYNOPSIS

Laminaria hyperborea has been, for a long time, one of the seaweed species of greatest industrial importance in Scotland arid other parts of the northeastern Atlantic.Also, much of the pioneer work on seaweed biolojr and chemistry was done on this species.Therefore, a review of the information available on the species was considered to be particularly valuable for all those concerned with seaweed research and utilization. This synopsis has been edited by G. Michanek, of the Marine Biology and Environment Branch, Fishery Resources Division, FAO. The author is extremely grateful to Dr. A. Jensen (Norway) who very kindly allowed her to use hie om drafts of the sections on harvesting, chemistry and utilization, thereby providing a more authoritative basis and saving her much time. Many people have been very helpful in providing information.For this the author is muoh indebted in particular to Dr. K. Ltning (Germany, FRG), Cand, real P. Svendeen (Norway), Mr. E. Booth, Dr. J,W.G, Lund and Mr, H.T. Powell (U.K.), Dr. A.H. Dizerbo (France), Professor J. Seoane-Camba (Spain), Dr, T. Hasegawa and Dr. S, Kawashima (Japan), Dr. G. Michanek (Sweden) and Cand, real 5. Sivertsen (Norway),She is also very grateful to the following for permisBion to quote unpUblished data:Cand, real P. Svendaen, Mr, A. Whittick (Canada), Dr. D.M. John (Ghana) and Cand, real O. Vahi (Norway).For permission to reproduce published figures she is indebted to Dr. K. LUning, Cambridge 1Jni,ersity Prese, Professor E. Baardseth (Norway), the Norwegian Institute of Seaweed Research and the Marine Biological Association of the United Kingdom, In the -text depths are expressed in metres below the lowest astronomical tide (LiT) which is defined as "the lowest level which can be predicted to occur under average meteorological conditions and under any combination of astronomical conditions" (U.K., Admiralty, 1963).Where necessary depth data have been corrected, usually -to the nearest metre, from the relevant tidal data, Where possible, algal names have followed the latest British check-list (Parke and Dixon, 1968) and animal names have followed the Marine Fauna of the Isle of Man (Bruce ,1963).

Distribution: Bibliographic entry: FAO Department of Fisheries Kam, J.M. (1971) FAO Regional Fisheries Officers FAO Fish.Synop., (87):pag,var. Regional Fisheries Councils and Commissions Synopsis of biological data on Laminaria Selector SM hyperborea Author ANE. Laminariaceae Taxonomy, morphology and anatomy.Distribution, ecology, metabolism and growth,Life cycle, reproduction, phenology. Population - structure, density and standing crop,Harvesting - -techniques, seasons,yielda. Chemical composition.Utilization.Selected bibi iography. FIRN/S87 Laminaria h,yrborea

CON T EN T S

Page No.

1 IDPNTIPY 1:1 1.1Nomenclature 1.11Valid scientific nazie 1.12Nomenclatural synonyme i 1.13Vernacular names

1.2Taxonomy i

1.21Affinities i 1,22Subspectee 4 1.23Genetic data 5

1.3Morphology and anatomy 5

1.31Erternal morphology 5 1.32Anatomy 7

2 DISTraBuTION, IXOLXY AND M1S OLISM 2:1

2.1Total area i 2,2Local vertical and horizontal distribution 2.3Effects of ecological determinants

2.4 Nutrition and growth 4

2.41Assimilation 4 2.42Respiration 5 2.43 Nutrition 5 2.44Growth 5

3 LIFE HISTORY 3:1 3.1Life Cycle 3.11Alternation of generations 3.12Nuclear phases

3.2Reproduction 1

3.3Phenology 7

3.31Seasonal variation in external appearance and morphology 7 3.32Seasonal variation in total fresh and dry weight 7 3,33Seasonal variation in other characteristios*

4 POPULATION 4:1 4.1Structure 4.11Age composition 4.12Weight or size composition 4.13Sporophyte...gametophyte and sex composition 8 iv FR14/387 LaninaHa hyperborsa

Page No. 4.2 Density 4:8 4.21 Avera«e densities of defined areas 8 4.22 Variation in density with ecological and accidental factors 8

4.3Mortality morbidity 11

4.4To1uantities (standing crop) of defined areas 12

4,5 Accompanying and competing species 14

5 HARV]ZPING 5:1

5.1 14ethod of Harvesting i

5.2Harvesting seasons 1

5.3Harvestin areas and depths 2

5.4 growth on harvested areas 2

5.5Total annual yields 2

6 PRcY1'ECTIO!r 6:1

6.1 Official regulaiions*

62 Control of ysicl features i

6.3 Control of chemical features*

6.31Water pollution* 6.32Artificial fertilization*

6.4 Control of bioloioal features i

6,41 Control of predation and competition 6.42Population manipulation*

6.5 Artificial culture i

7 CHII4ICAL COMPOSITION 7:1

7.1 Water content

7.2Inorganic constituents 2

7.3Organic constituents 4 7.31 Carbohydrates 4 7.32Proteins 6 7,33 Fat 6 7.34Pigments 6 7.35 Other organic constituents 6 FI1'1/sO7 Laminaria hyperborea y ?ae No.

B UTILIZATION

8.1Human food 8.2Animal fodder 8,3 Manure 8.4Industrial products

9 REVERENCFS 91

* AB no information was available to the author these items have been omitted from -the tex-t, FIF/S87 Laminaria hyperborea 1:1

1 IDEl'ITY ChinI (1968) g-ives 22 names for the species, most of which are in very local uno if at all. 1.1Nomenclature

1.11Valid scientific name 1.2Taxonomy Laminaria hyporborea (Gunnerus) Foslie. 1,21Affinities This was described and figured by Gunnerus - Supreneric (1766) as Focus hyperboreus, but recombined as Laminaria herborea by F'oslie (1885). Laminaria hyperborea (Gum-io) Foal, belongs to the family Laminariaceac of the order 1.12Nomenclatural. synonyme Laniinariaies of -the division Phaeophy-ta. Many synonyms are given by Foslie (1885), -, Generic De-Toni (1895), Dii Rietz (1920, 1953) and others, the most important being the following: The genus Laminaria was set up by Laxnourouz (1813) with L, digitata as the -typo species. Fucus scoparius S±rm, 1762 The following in a dia-iosis of the genus. Focus cijjtatus Nohr, 1786 Laminaria digitata tyngbye, 1819 The sporophyte is in three distinct parts, Laminaria phycodendron la Pylale, 1824 holdfast, stipe and frond (lamina or blade), Hafgygia digitata KUtzing, 1843 The boldfas-t may be a disc or more commonly Laminaria cloustoni Edniondston, 1845 formed of branched haptera which in a few species Hafgygia cloustoni Areschoug, 1883 give rime io more than one stipe.The Ertipe la Laminaria digitata f. typica Foslie 1884 simple (occasionally branched following damage), terete or flattened, and expanded into a smooth The earlier naine Fucus acoparius of frond which may be entìre or spi1t into segments Strm (1762) was invalid because, although it distally,The primary growth region is the tran- clearly applied only to this species, it was sition zone between stipe and frond,The frond published in a work not generally employing has no midrib but may be elevated or depressed binary nomenclature (Du Riets, 1953). in different areas,It has no cryptoetomata or natural perforations,Mucilage ducts may be For many years there was confusion between absent from the plant or present in the frond this species and. L. digitata (HW1S.) Lamour., and sometimes also in the atipe.Unilocular e.g. by Lyngbye (1819), and Harvey (1846-51) and sporangia are borne in irregular dark--coloured although Le Jolis (1855) clearly distinguished sort on the frond.The zoospores develop into the two species and listed their reliable microscopic sexual dioecious garnetophyteseach characters some confusion continued and even female unicellular or a simple or branched Foslie (1884, 1885) misidentified some filaxent, each male rnu].tioeiiuiar but varying specimens (.1iuida, 1965).He (Foslie, 1885) was also in complexity, satisfied, however, that Gunnerus (1766) had not included L. digitata in him description The genus is usually divided into two sec- because he examined herbarium inclu- ibas, the Simplices with an undivided frond, ding the type specimen,Unfortunately Le Jolis the Digitatae with the frond normally split into (1855) had not been of -this opinion and thought segments.Well over a hundred species of that Gunnenus' description was thus invalid, Laminaria have been described but the majority and thai Edmondaton' s L, cloustoni was there- of these naines are inmalid, having been trane-- fore the first valid name,This synonym was ferred to other genera or being synonyms of used for many years, particularly in Britain valid specific names,There romains considerable (e.g. Newton, 1931). confusion about many of -the specific naines still in use,Thim is mainly because many species are 1.13 Vernacular naines rnorphologically plastic and only recently have serious attempts been made to define specific Norway:Trolitare, sortare, s-tokictare, palme- bouncIHes by experimentation.There is confu- tare, akramehes-tatare, havtare, kurvtare, sion both within small geographical r 'cas whore stolpetare(Grenager, 1953). perhnim a few species show many form.'rroncousl.y termeL species and between widely separated areas British Isles: Tangle, redware, cuvie, where perhaps the same specific epithet is used cuvy. for d.ffering entities,In view of the taxonomic chaos in which the genus mua-t be considered to be, Germany:Palmentang. -the following lis-t must be considered as a guide only.Much more research must precede a defin1- B'rance (Brittany):Tall-penn, tali-ebrel. -tive list of species.The Simplicee section is discuesed in more detail by Burrows (in prepara'-' tion). 1:2 FIHM/S87 Laintharia hyperborea

Species of the genus Laminaria ai-e listed L. cimoifolia J. Ag. Doubtful species.Accox- below in approximate order of geographical dis- ding to Wilce (1959) this naine is invalid tribution, starting with those occurring in the and the plants attributed to it indistin- Northeast Atlantic and progressing westward, guishable from L, .enlandica (Wilce, 1965) but Druehi (19.:retains the species. L. hyperborea (Gunn.) Fosl, Distinct species. Northwest Atlantic, Northeast Pacific. Northeast Atlantic. L. platyineris la Pylaie.Doubtful species. L. digitata (Huds,) Laisour,Distinct species This is variously regarded as digitata or with several varieties or forms induoed by simple and included in L. groenlandica by habitat.Northeast and Northwest Atlantic, Druehi (1968) and in L, digitata by South Arctic. and Cardinal (1970),Northwest Atlantic, Northeast and Northwest Pacific, L.unneri bsl.Indistinct spcies of local occurrence.North Norway. L, fiasilis J. Ag,Doubtful species,Northeast Pacific. L. ochroleuca la Pylaie,Distinct species. Northeast Atlantic. L, bongardiana Post. et Thipr.Doubtful species. Northeast Pacific. L. pallida (Gravo) 3. Ag0Distinct species. Southeast Atlantic, Southezn Ocean0 L. sinclairii (Nary.) Burl.Distinct species. Northeast Pacific. L. schinzil Posi.Somewhat doubtful species, of local oocurrenoe,South Africa, L. ephemera Setch.Distinct species.North- east Pacific, L. rodrjuezjj Bornet.Distinct species0 Mediterranean. L. farlowil Setchell,Possibly a distinct species but of local occurrence.Northe L. saocharina (L.) Lainour,This and the next Pacific. two species are only distinguishable by the presence or absence of mucilage ducts in L. complanata (Se-tch. et Gardn.) Setchell, the stipe and frond0Wilce (1965) Probably a diErtinc-b species.Northeast aggregates them as L, eaccharinbut as at Pacific, least one is genetically distinct (Burrows, 1964) the species delimitation L. setchellil Silva.Distinct species. North-- is uncertain.Northeast and. Northwest east Pacific, Atlantic, Arctic, Northeast and possibly Northwest Pacific. L. rwrechtii (Areseh.) de Toni.Local occurrence,Alaska, L, groenlandica Rosenv,See notes on L, saccharina.Arctic, Northwest Atlantic, L. dentigera Kjellm,Possibly a distinct species. Northeast Pacific. Northeast and Northwest Pacific. L. agardhii Kjellm,See not os on L. saccharina, L, lontipes Bory,Distinct species,Northeast Northwest Atlantic. and Northwest Pacific. L. atrofulva J, Ag.Doubtful species of local L. oensis Niyabe,Variable species. North-- occurrence,West Greenland, east and Northwest Pacific, L. nigripe J. Ag.Thought to be distinct by L. ensiformis J. Ag. Doibtful species of local some authors bu-b Wiloe (1965) suggested occurrence,Northwest Pacific. synonymy with L. diibata.Arctics North- west Atlantic. L. ponica Aveach.This and the following three species were shown by tabu (1964) to L, longicruris la PylaieProbably o dirrtinot he interfertile,Some are eophioally species though -there has been doubt about separated and may be scads (Ilasegawa, 1966). mucilage dueto,No'-bhwest Mlentio. However, as transplantation experimente have not been reported it is not knownt L. faeroenais Borgesen.Doubtful species because present whether they are etioally its difference from L. lonj.orurìs is distinct. based ou iauoilao dueto about which there is uncertainty.Northeast Mlantio. L, religiosa Miyabe.See notes on L. ,ponica, Northwest Poific, L. solidunciic J, Ag. Distinct species,North- west Atla'ttio, FIr4/S87 Laminaria hyporborea 1:3 L. ochotensis Miyabe.See noten on L. japonica. Note on the benzicline tee-t: Northwos-t Pacific. Two solutions are prepared; A. 5 g bee zidine, L. diabolica Miyabe.See notes on L. japoaica. 25 ml 25 percent HC1, to 1 000 ml with H20; Northwest Pacific. B. 10 g NaNO2, to 100 ml with H20,Equal quantities of A and B are mixed five minutes before L. fragilie Niyabe.Doubtful species of local use and used for up to two hours,Freshly cut occurrence.Japan. tissue of L. hyperborea (and of some other algae) turns bright red immediately on application. L. longipedalis Okam. Somewhat doubtful That of L. digitata changes to yellow-brown species of local occurrence included in slowly. L. d.iabolica by Niyabe and Nagai (1933) CAIJTIC:Users should be well protected from and separated by 1liyabe (1902, 1936). bensidine because it is highly carcinogenic and Japan. can pass through skin, L, cichorjoides Miyabe.Distinct species. Specimens of L. hrperborea are deposited Northwest Pacific. at the British Museum (Natural History), Cromwell Road, London 5.11,7, L. anustata Kjellm.Distinct species.North- west Pacific. The following key has to rely largely on the presence or absence of mucilage canals as L. coriacea Miyabe,Distinct species.North- distinguishing characters,These may not be west Pacific. very reliable and are often difficult -to see. Many of the species of Laminaria must be treated L. sachalinensis Miyabe,Probably a distinct with caution, as explained earlier.All the species,Northwest Pacific. very doubtful species and those of very local occurrence are omitted. L. siko-tanensis Miyabe et Nagal.Local ooc'.x'- rence,Northwest Pacific. 1. Frond normally digitate (splits may fail to appear in very calm water) 2 L. yendoana Miyabe.Probably a distinct species but of very local occurrence. Frond entire 11 Japan. 2, Holdfasi a disc 3 L, subsirnplex Miyabe et Nagai.Doubtful Holdfast formed of haptera species included in L. cxuneifolia by 5 Setche].l and Gardner (1925) but retained No mucilage ducts in frond by Nagai (1940).Northwest Pacific. or stipe, sonelongated t, ephemera L, taeniata Post. et Thzpr,Doubtful species, Mucilage ducts in frond but not stipe 4 Northwest Pacific. Stipe terete but thicker near L. ualmaeformiE Olcam.Probably a distinct the base, warby outgrowths on species.Northwest Pacific. stipe and holdfaat when oldL. yezoennia Stipe flattened distally, L, gurjanovae A. Zin,Distinct species recently lamina deeply split L. palmaefomia described.Northwest Pacific. 5, Stipe torete but thicker near the - Specific base, tapering just above hap-tera, with mucilage ducts 6 herbarium ja in Trondheim Museum but the type specimen of Laminaria hyperborea Stipe cylindrical at base, torete or appears to be lost. flattened distally, with or without mucilage ducts 8 Diagnosis of L, hyperborea: 6. Stipe rugose except when young, Stipe terete, thicker near the base, rugose rigid, mucile duc-ta present except in young planta or new growth, rigid, L.,yperboroa containing mucilage ducts.Very young stipes bipe smooth 7 of 1-20 cm distinguishable from L. digitata by a narrowing at the transition zone.Frond 7, Prend dark, occurs in Pacific developing from November onward separated from L. dentigera old frond by a narrow collar after the first Stipe and frond lit in year.Sortis covering most of old frond for 1-2 colour, occurs in Northwest months between September and April.Tissue Atlantic L, ochroleuoa positive to benzidine -teat (Jensen and 11aug, 1952). F1fTh1/387 Laminax-a hyperbqz-a Stipe light in oolour, terete, 21.No mucilage duc-ta in a-tipe L. saccharina occurs in South Atlantic L. pallida Mucilage ducts present in Stipe dark, may be flattened stipe L. groenlandica distally, smooth or rilgose 9 22,Stipe 2-3 mm in diameter, Stipe smooth and flexuose, devoid mucilage ducts in frond but of mucilage ducts which are not stipe L. gurjanovae present in frond L. digitata Stipe 4 nie or more in diameter 23 10 Mucile ducts present in stipe 23.Stipe of mature plant 300-600 nia Mucilage duc-ta near surface of long, mucilage ducts in both stipe, stipe nearly black, smooth a-tipe and frond L. lonipedalia flattened distally L, nigripes Stipe less than 300 mm long 24 Mucilage ducts in mid-cortex 24,Median fascia less than a of stipe, stipe smooth, flat- quarter of the width of the tened distally L. setchellil frond L. angus-tata Stipe hollow L. longicruria Median fascia more than a quarter of the width of the frond 25 Stipe solid 12 25.Margina of the frond crispa-te Holdfast a disc 13 and waved throughout its length, Holdfast of haptera 15 mucilage ducts in both stipe and frond t. oichonioiclea No mucilage ducts in either frond Margina of the frond not orispate, or stipe, sari elongated L. ephemera and not waved near the transition Mucilage ducts in frond 14 zone 26 No mucilage ducts in stipe, 26,Median fascia more than three- stipe thicker near base, fifths of the width of the frond, Pacific L. yezoenais coniaceous and rigid, mucilage Mucilage ducts in a-tipe, son ducts in frond but not stipe L. coriacea rounded, Northwest AtlanticL. soli&ungula Median fascia less than three- fifths of the width of the frond 27 Hap-tera giving rise -to more -than one stipe (rhizoma-boua) 16 27.Mucilage ducts in frond but not a-tipe L. saccbunina Single stipe per holdfast 18 Nucilage ductsinboth frond 16,Mucilage ducts present in frond, and stipe 28 absent from stipe L. longipes 28,Eiullae present on median fascia Mucilage ducts present in frond except in very old fronds and stipe 17 haptera vertici].late L. sachalinensis 17.Stipe smooth, occurring in Bullae only present in very Mediterranean L. rodriguezii young fronds L. japonica group Stipe rugose, occurring in Northeast Pacific L, sinclainli 1,22Subspecies There are no subspecies of L. hyperborea, 18,Stipe flat for dia-tal two- ilowever, -two forms other than f. h,yperborea thirds of its length L. complanata have been described.The first was by Fbslie Stipe mainly tenete 19 (1885) and was L. hyperborea f. compressa Fosli.This was a renaming of his L. digitata Occurring in Atlantic, Arctic, f. lon'-ifolia Fblie (Fkalie, 1884).The form Northeast Pacific or Bering Sea 20 is figured only as a transverso section of the a-tipe, which is flattened,As no specimen of Occurring in Northwest Pacific only 22 this furm can be traced in Fosli&a herbarium and tbstrong possibility that this was a form No mucilage ducts in either frond of L. iligitata cannot be eliminated, it would or stipe L. agandhii seem prudent toigoore the description at Mucilage ducts in frond 21 present. FI14/S87 Laminaria hyperborea The second forma is L0 hyperborea f. attacked by the limpet Fatinpellucida (L.) the cucullata Svendsan ei Kamand is an ecad charao- haptera produced nearest to the cavity formed teristic of very calm water (E3vendson and Kam, are clearly stimulated in their growth and 1971).It is found in vary sheltered areas and branching so there is a tendency for the cavity in comparatively deep water in slightly exposed to be covered up and extra attachment to the areas, mainly in Norway.It is distinguishable rock formed, from L, jaia f. cucullata Le Jolis which it closely resembles by the taper near the base of The stipe is terete for the whole of its the stipe, the collar between the new and old length, except where it flattens suddenly into frond and the positive reaction -to the bensidine the transition zone distally,Ii tapers slightly test. over most of i-ta length bu-t mainly near the base and apex,The maturetipe is sufficiently 1,23Genetic data rigid to bear -the weight of -the frond out of water with little bending except near the tip The chronic some number in young sporophrtea, but can usually be forcibly bent into a complete unilocular sporangia and female gametophytes circle before snapping and showing a clean break, has been counted by Evans (1965).Because of cept when newly formed, the surface of -the the small and mixed size of the chromosomes stipe is rugose or papillate and is thus a the counts were variable but he thought the suitable substratum for many epiphy-tes. most likely numbers -to be 31 haploid and 62 diploid, At the transition zone the stipe expands into the frond,The frond is a flat and very In the north Ehropean species of Laminaria smooth blade normally divided into fingers,The it seems that the species boundary coincides shape of the- base depends on the habitat b-ut is with an interfertility barrier.This conclusion usually truncate or slightly cordate. is made from Schreiber's (1930) beautiful set of experiments in which he failed to obtain The morphology of the plant is markedly hybrids between L. hyporborea, L. digitata and affected by water movement.Where there -la L, saccharina,On the other hands Sundene (1958) strong wave action the holdfast is well developed and Svendaen and Kam(1971) obtained normal with thick, well branched haptera and a large sporophytes from crosses of gametophy-tes from area of attachment.The stipe is fairly rigid different forms or ecads of L. digitata arid and more or loss straight.The frond is L, hyperhorea respectively. leathery, truncate at the base, fan shaped and. divided into many segments, 30 fingers being Chapman (1948) suggested that differences common.The outer fingers of the new frond in the vertical distribution of L. hyperborea (Fig. lB) are released early (February) and the with latitude might be attributable to different old frond is often lost in March,With decrea- strains of the species. Kam (1969) grew plants sing exposure the number .of fingers of the frond from spores from Norway and the Isle of Nan at decreases and -the old frond remains attached different irradiances and temperatures and untjl later.Where there is a strong current observed almost identical growth.A more likely but little wave action, the holdfact is also explanation for Chapman's observation is his well developed bu-t the stipe probably more alternative suggestion thai ii is due to dif- flexible,The frond is cuneate at the base and ferences in ecological conditions. fingered but much more elongated -than the fan- shaped wave action form.The plant is then 1.3 Norphcloy and anatoniy similar -to L. digitata under similar conditions, particularly in the shape of -the frond. 1.31Erternal morphoior Ii-, conditions of shelter from both wave The mature plant is distinctive and well action and water currents, the plant has a very differentiated (Fig. i).The holdfast consista different appearance and is termed f. cucullata, of haptera which have grown out of the base of The holdfast is poorly developed with few and the stipe.The stipe base is in the shape of weak haptera and weak adhesion to the rock, an inverted cone, the lower tip being the The stipe is bent or tortuous and short, nor- point of original attachment but which may not mally being lesa than 50 cm.The frond is thin, still be in contact with the rock.Around this brittle and easily torn, corda-te athe base, position there is a ring of anali haptera and entire and broad, forming a drapingapron5 progressively more recent hap-tera produced (Svendaen and Kam, 1971). upward from here are arranged in about seven to eight vertical rows, each of up to about ten The frond: stipe weight ratio at any g-von haptera. ch hap-teron is terete, up -to about age increases with shelter and with depth 10 mm in diameters and is branched about four (Kam, 1971m). -times, the branches decreasing in diameter. The -tmpa of the hantera are expanded into small For -the maximum size attained by this diccn adhering to the rock.When a holdfant is species see 4.12. 1:6 FIi4/S67 Laminaria_hyperborea

Frond (Lamina) (Blade)

Fig, IDrawings of sporophyes of L. hyperborea A, Whole plant during the slow growing B. Part of a younger plant in February eason. showing old and new fronds. FII/S87 Laminaria hy-erborea 1:1 1 32 Anatomy continues on the outside, the spaces appear

deeper in the cortex and enlarged, with oece-- Killian (1911) descrIbed the anaton' of L. atonal connexions to the meristodexis,At the ditata in detail and compared i-t with that of inner margin, groups of small cells appear and L. hy-oerborea, divide,They form an irregular layer on the inner surface of the ducts have dense contents The mature e-tipe is differentiated into a and, are secretory,Secretion of mue-linge number of different tissues,Some originate appears to originate from the Golgi vesicles from prïmary grow-th and some from Becondary, (Schnepf, 1963).The epidermis of the maturer The transitjon zone is responsible for primary part of the stipe, well below -the transition growth,The activity of this zone results in zone, where the surface is rugose, cOnsists o? elongation o' the stipe below and production of' a brown, fairly thick, corklike layer (Schultz-. the frond above.,The meristematic tissue is Schult zenstein, 1853), maid (e.g. by Killian, 1911) to be confined to the surface layer of celle 1m-t these could not Secondary growth in thickness is accomplished. by themselves account for ail the growth in through a secondary merle-tern situated in the length so it seems likely that the inner Lineuù cortex inside the mucilage ducts (Fig, 2A). How mue-t also divide,The surface meristem, or the tissues exterual to the merim-tem are meristoderm, consists of small cells containing replenished as the circumference increases, is chromatophores and covered by a layer of mucilage. not olear bu1 the meris-toderm may be reeponeible. Radial rows of similar cells are cut off centri- The secondary meristem produces cells of the pa-tally.Passing inward. (Fig, 2A) these outer cortex only.The medulla of the stipe gradually become wider, elongated and paler in thus remains -the saine diameter as is formed by the outer cortex.The cells have pointed ends primary growth and is thus wider dim-tally, but and are arranged less regularly.Cells of the the cortex increases annually,During the famt inner cortex, in-to which -theme grade, are longer growing season each year, an approximate cylinder still, have blunt end.m and walls thickened with of cortical tissue is tima laid down outside the mucilage,Some of them have small tubelike stipe tissue of the previous year.Thiring the outgrow-the.The cells of the inner and outer slow growing period few cells are laid down and cortex show up in different colours under a these are darker in colour then fas-t grown cells. polarization microscope (Baardseth, 1964). This is apparent to -the naked eye but hardly Passing inward the cell rows become more distinguishable microscopically,It results Lin separated from each other by mucilage but in annual growth rings (in transverse aectidn) or some cases are still joined horizontally by lines (in longitudinal section) (Kath, 1963), their original pit connexions.,In L, digitata The dark line for each year ham -the same length and Saccorhizaolyschides (Killian, 1911; as the stipe at -the time of the slow growing Sauvageau, 1918), and therefore probably in season during which it was formed,Figure 3A L. erborea, more numerous horizontal con- shows diagrammatically the slow growth lines of nexions between these filaments are formed by a a twoyear old plant, the stipe of which was process with the partial appearance of conju- 100 mm long a-t -the end of -the first season of gation.Opposing papulosa outgrow-ths on fast growth and 330 mm long at the end of the adjacent filaments grow toward each other and second. join, With a breakdown of -the separating walls but no fusion of nuclei,During growth in length Haptera are produced during the fast growing -the medullary cells appear only -to be produced season,The extezal cell layers of the stipe successively from the inner cortex and they divide tangentially and produce a hump In which consequently become stretched.As the septa of there is an apical merietem (Killiazi, 1911), the original longitudinal cells seem fairly Growth in length continues until the hapteron rigid the cell walls in these regions are pulled reaches the rock surface when it expands into an in-to a trmmpet shape,The swollen ends are irregular disc,There ici no medullaina hap- filled with granules,A-t the same timethe teron but the interual cells are elongated, cross connexions lengthen and become mainly Thicilage ducts are present,The epidermal cells oblique.In addition to these two types of have few chroma-tophores at first, so -the hap- cells in the medulla, there are later formed the toron is palo, bu-t moro later uhcn the hapteron socalled hyphae as outgrowths from cortical darkens, Numerous rhizoide grotT out rrom the cells (Fig. 2A).These become very numerous disc and become attached,If one haoron ring and grow mainly horizontally through the muci- is formed during each fast growing period, then laginous spaces between the other, thicker, each hapteron is separated from -the next by a rnedullary elements,The medulla thus consists slow growth line.Thus tracing each line to the of filiform ramified tubes entangled into a outside between the haptera demonstrati's the felt. number of hap-tern rings formed each yearwhich varies,The shape of the base of the sipe Mucilage ducts of the stipe arise (Fig. 3B) am an inverted cone is exolairud by below the transition zone as lenticular -the increase in the thickness of' the Stipe with spaces between the outermost cells in the outer each year of secondary g-row-th which eitcndn oniy cortex (Guignard, 1892). A cell division down to the last ring of haptera, 1eaving thc 18 FI14/s87 Laminax'ta hyoerborea

Mucilage Secondary lOOpm duct meristem

Mucilage ducts

gig. 2Anatomical drawings of L. hyperuorea Half a longitudinal section of a young C. Physodea in medullary (above) and ipe, surface (below) cello (magnification about four times that of A and B), 'l'ranoverse section of frond (me scale). from rdaeth(1958)., FIrA/S8 Lnminaria orborea

0 0000 0 00 o o 00 sS7 B cS6 A : :::: ::: : :o;0,0::, :°:::. 0000 0000 00 00J : : 0000000 °° f S3 ,k

20mm

sS2

C E F f S2

cS2 sSi

Is'

Pig. 3 Diagrenmatic representation of slow growth lines in the stipe of L. hye, orea A, Median transverso section of whole stipe of CF, Similar sections of 2year aids sanpied 2year old, during the fast growing aeason.Primary B. Median transverse section of the base of tissue shown black, 7,rear old. £51, f52, f53, fast secondary growth of ist, 2ndand 3rd years respectively;oSi, n52, 1353, slo secondary growth of lot, 2nd and 3rd yearn peotivel.y. 1lo FI}1/B8Laminaria hyperborea stipe below only as -thick as that of -the The frond is also produced by -the activity previous year. of the transition zone.It has basically the same structure as -the a-tipe (Fig. 2B), and the PlaMe oen be aged by observation of -the meristem, cortex and medulla (but not -the slow grow-th lines (Kam, 1963).After removing mucilage ducts) of the -two structures are can- all bu-t a few millime-tres of each hapteron, -binuous with each other.Jus-t above -the transi- the base of -the e-tipe is cut long-i-tudinally tion zone the lamina is -thick but lateral and centrally, passing -through a row of hap-bera. expansion, partly due to anticlinal division of A thick section is then -taken,In Figure 3B the ineris-toderm, results finally in a thin blade which is diagrammatic only9 -the primary bissue with a thinner epidox-mal region than the stipe, is shown black al-though in -the plan-t -the medulla a limited number of cortical cell layers and a is more easily distinguished in -this region. confused network of medullary elements lacking Near the base -the junction of the primary and the mainly longitudinal arrangement of the e-tipe. secondary tissues can appear slightly darker There are fewer hyphae than in the e-tipe.There and be confused with a slow grow-th line,If is some doubt as to whether the medulla cells -this turns centripe-bally when followed down -to arise only from the transition zone or also from the base it is the primary/secondary tissue cortical cells of -the frond,The lateral stret- junction, bu-t if it -turns outward over the firat ching results in the cross connexions in the a-t-tachmen-b disc or haptera -then i1 is a slow medulla becoming longer -than in the stipe and growth line dating from when -the plant was very more cell walls forming across them.When the small.La-ter lines can be counted more easily frond expands and becomes thinner, both medulla (Fig. 3B), particularly if -the separation by and cortex are reduced bu-t the cortex is less haptera is noted.A count should be made on reduced than -the medulla, both sides.If the plan-t is sampled during the faa-b growin, season from December to June, care Mucilage ducts in the blade are produced mue-t be takiin -to determine -that the last slow just above the -transition zone (Guigeard, 1892) growth line has been made apparent by -the and are positioned in the cortex as in the stipe current fasi growth.If, as in Figure 3C, the (Fig. 2B),They are spread ou-t in an anas-tomo- las-t apparent growth line was inside -the ou-ter sing network beneath each surface of the frond, ring of haptera which were clearly dark and with no connexion between -the two systems. rugose and therefore a year old, -then -there must be another slow growth line not yet visible but The, splits in the frond of L. hyperborea which should be counted.If, on the other hand, are no-t accidental tears but are preformed by as in Figure 3D-there are no haptera on -the the activity of the -tissues (Killian, 1911), outer side of -the last line, -then secondary Longitudinal furrows appear in the epidermis on thickening of the stipe has already started and one or both sides of the frond, these deepen and demonstrated the las-t line, though haptera have the internal tissues rupture.The epidermis then no1 yet developed.These can be seen -to be growe over the edge of -the cortex and medu12. developing in Figure 3E.Finally, when new hap-tera have developed, as in Figure 3F, the In common with other Fhaeophyta, L. hyper- line is again inside a hapteron ring but these borea cells contain vesicles of f'ucosan, or hap-tera can be easily distinguished from -those physodes. - The appearance of these in the surface a year old because they are pale and smooth and cells and in the medullary elements is shown in may not have expanded at their i;ipe.Infestation Figuro 2C (Baardee-bb, 19&).The latter are by Patina pellucida makes age determination apparently more distinctive of the species. difficult firstly because much of the tissue is removed and secondly because "interference" lines and additional hap-bera are produced as a reaction to -the infes-ba-bion, FIT1/S87 LamjJyperborea 2:1

2 DISTRIBUTION, XOIOGY AN]J MFYrABOLISM the Devon (southwest Ebgiand) coast to turbidity, the lower limit near Dartmouth being 5 to 6in 2.1Total area and that off Stoke Point, in clearer water, 17 s Another factor which can affect the lower limit L. rea is conLned to the northeast is grazing (Jones and Kam, 1967). Atlantic (Fig. 43TIt was reported absent from Spitzberen b' 5vendsm (1959) but more recently Near the upper limit in exposed sites the Dizerho (1969) has reporLcd it present from plants are no-t normally the larges-t found in specimens collected by Noign (1965).,These have the area, presumably because of removal by wave not boon checked with the benzidine test and are action.The best developed forest is thus a few not typical of L. hyperborea from farther south. metres below the tipper limit when the vertical There is little available information on range allows,With increasing depth the density Jan Mayen or Bear Isiand,Before Dizerbo'a decreases and -the maximum size of the plants may Spitzbergen report the most northern known occur-. get smaller,In some areas, e.g. the Isle of renco was the North Cape in Norway and the plant Man, the lower liai-t is determined by fac-tors extends westward along the north Norwegian coast (such as grazing) acting on the establishment and a short distance into the U.S.S.R. of -the species rather than its later growth rate (Vozzhinskaya, personal communication).It also (Kam, 1963).In this cases the deepest plants extends south along the rest of the Norwegian are relatively large and grow only slightly more coast and along the west coast of Sweden into slowly than shallow plants,In other cases the Ka-ttega-t (Roeenvinge and Lund, 1947).Accor- e.g. in southwest Norway (Kam, 197m), the ding to J6nsson (1912) it occurs on most of the deepest planta grow very slcwly and are much coast of Tcland except part of the east, being small er, present onl:r at the mouth of Bermfj73xur.How- ever, Monda (1969) has reported i-t from 2.3E'fects of ecoloçical determinan-ta Reyarfjrar, further north into this eastern region affected by the Arctic current,It This species is clearly unable to with- occurs in the Faeroes (Edaondston, 1845), the stand prolonged. exposure to the air,Its

Orkneys (Spence, 1918), around most of the pene-bza-tion into the intertidal zone is nor--

British Isles (including the Scilly Isles, mally confined to eonsïderably lesa than a ver-- Harvey, 1969) and around the Channel Islands tical metre above LilT or to rock pools where it (Kam, 1961),Ii is present in Helgoland. but in totally immersed.Kitching (1941) nt-tribu-toe absent from the rest of Germany,There is none itsiack of success in the intertidal compared on the Dutch coast (Hartog1959)oItin with -that of L, jata to -the greater exposure present on the coast of Normandy (Le Jolis1855) of the frond to desiccationbeing held. up by a and Bri-ttany (Sauvageau, 1918) and southward s-tiff erect stipe,Ulning (unpublished) has (Hamel, 1939; Hoek and Douze, 1967) observed mass mortality of L. hyperborea, with but rare on the Basque (Feldrnann and Lami, 1941) L, digitata surviving, after unusually low sea coasts of France,I-t reappears on the north levela-t Helgoland in winter,This may have coast of Spain where the -temperature is lower been due to a lack of low temperature tolerance., (Hoek and Bonze, 1966).According -to Seosne The relatively inflexible stipe may also he Cainba (1966i-ta southern limit is in Portuga]. more vu]iterable to damage by breaking waves and a-t about 40 20 N. cause more leverage on the holdfast than that of L. digitata. 2,2Local vertical and horizontal dis-tribu-tien L. hyperborea grows only on solid rock or s-tones or boulders large enough not to be turned The upper limit of L, Iryperhorea is over frequently by storms,On solid rock it usually just above LAT (lowest astronomical grows on horizontal or upward facing sloping tide) (Kitching, 1941;Icain, 1962) or depressed surfaces.It is rare on vertical faces or below -this under conditions of extreme exposure slight overhangs and then confined to small 10 wave action (Kam, 1971a), by reduced ledges.I-t is absent, from completely downward salinity (Sundene, 1953) or occasional other facing surfaces,On large boulders with flat factors,The lower limit is much more variable tops and steep sides the edges form a more and depends on local factors,Light is the favourable substratum than the central -top under most obvious of these and in very turbid water forest forming conditions because olesa shad- such as the Menai S-traits off North Wales, ing there by other plants,. When no.i1iy stable the lower limit is only 2,m below LAP boulders are turned over by unusually severe (Knight-Jones et al,, 1957) and in some polluted s-bornsplants of L. h,yperborea with their areas the vertical range may be reduced -toIii stipes horizontal may survive for some time but (Bellaniy e-t al., in press),In clear water, -the stipes bend to -the vertical during growth.,, however, the species can he found at more than 30 in below LAP, e.g. a-t 34 in in Norway (Kam, There is no direct information on the 1971a) and 36 m in southwest flgland (Whittick, -temperature tolórance of mature planta of thiu 1969),Fors-ter (1955) attributed variation on species -though there is come mnforma:tion on the Rig. 4The geo:phical digtribution of L. hyperborea.The arrow indicajee 1he uncertain Spitzbergen record. PIR58 Lanaa Aorborea early stages.it is likely that the optiîmun order as the minimum requirement of garnetophytes temperature (given adequate light) for gasse- and microscopic aporopb,y-tes0In the summers how tophy-te and young sporophyte development ertena ever, these stages, but not later sporophy-tes, from 100 to 17°C (Kam, 1964, 1965).Above 17 C could well be saturated at this level,At the surv-ival of gametophyïes is progressively shallower depths if there is no shadings all reduced, few surrivinat 20°C, while sorophytes plants could be saturated in summer down -to 5 n are produced up to 19but not at 20°C (Kam, but almost all would be light limited in winters 1964, .1969).However, sporophy-tes can grow at though growth would be possible except where 20°C in the laboratory (Kam, 1965).In shading is severe.These estimates are supported Portugal, at the southern limit of the species, by observations of growth a-t different tines of the winter surface sea tempera-turo during year a-t these dePths.In Helgoland (tUning, spori-ig drops to about 14 C so presumably young 1971), even at 2.1 m below LAT the irradiance stages could survive initially further south. does not rise above the compensation point for However, in this region the maximum summer fronds until April and only in June and July is -bemperatuz'e,which has -to be tolerated by later it above saturation,This is attributed mainly sj,orophytes and. maturé plants, is 19° to 20°C -to ses-ton stirred up by storms in winter,It is (U.S. Navy, Hydrographic Office, 1944).It thü likely -that off the southwest coast of Norway eeemalikely that later stages have an upper (Kam, 1971a) light penetration through the sea temperature tolerance similar -to the early stages.is better in dn-ter than off the Isle of Man, Less is known about -thé tolerance to low 1en- Irradiance levels in deeper water may thus be peratures.Young sporophy-tes grow slightly higher.In addition to transmittance through the more slowly a-t 5o) than 10°C (Kam, 1969) and water shading is a Very important factor,Small can thus presumably survive lower temperatures. algae, silt, etc. can cause severe shading on the The surface sea temperature on the open coast.. rock surface and mus-L reduce considerably -the in north Nc rway drops to about 3°C in winter irradjance important for the growth of the very (Saelen, 150).On the west coast of Spitzbergen early s-Lages, even in a fairly open community. in summer (July to September) the surface tem- The Laminaria forest itself, of courses is far perature records vary from 20 to 6°C (Heiland- from being open and, also causes considerable Hansen and Narisen, 1912;Nansen, 1915; reduction in irradiance,Hitching (1941) found Svendéen, 1959).1t is not clear whether the -that -thick L, hyperborea forest in Scotland cut northern uni-t of the species is determined by out 98,8 to 99.1 percent of the light,On -the winter temperature or the lack of light during other hand, in preliminary measurements in the the obligate winter sporing season0 Isle.of Ma(Hain, unpublished) i-t was foüM that this species cut ou-t only about 75 percent of th The quantitative light requirements of the irradiance, early stages of L, hyperborea are known.Zoo- spores can undergo their first development in L. thrives in moderate water the dark and some can survive as early gameto- movevient.The stipe has enough flexibility to phytes in darkness for at least 80 days at bend to and fra with the horizontal movement of 10°C (iain, 1969).A minimum of about wave action on the bot-tom subiidally,It cannot 2 pg. cal/cm2 seo (or 20 lux of daylight) is withstand severe breaking waveshowever,Its necessary for the development of the gmetophytes place at the top of the subtidal zone is then whch are light satura-ted at about 60 ig. calf taken by Alaria esculenta (Kan, 1971e).It cmsec at 10°C but higher at 17°C (Kam, 1963). thrives also in sites erpsed to strong currents, Very early sporophytes have light requirements for example at the entrances to sea lochs and similar to gaine-tophytes (Kam, 1969) but both fjords.Where there is lit-tie or no water move- the compensation and saturation levels increase ment the plants develop into f, cuoullata, with increase in size0Below saturation the growth in light and dark periods is a product Little is known of -the salinity tolerance of the light period and -the irradiance, e.g. of the species,In an experiment on the early 40 xg. cal/cmsec for 12 hours in 24 would stes, Hopkin (unpublished) observed very give the saxe growth rate as 20 mg. cal/cm2 sec occasional and slow sporophy-Le development of continuous irra&iance (Kam, 1964).The following gasnetophyte maturation at 16°16o5 but requirements of intermediate phases are not not below this.It is almost certain th.a-t its known bu-t in mature plants the compensation absence from the Baltic is determined by the pont for photosynthesis is about 60 ng, call reduction in salinity.It occurs in the outer cm sec and saturation is at 300-500 ,ug. caif part of the Oslofjord only from 8 to 25 mits cm2 ecc (LIming, 1971).The amount of light absence from the shallow subtidal being ait ri- reaching the habitat depends on surface buted to reduced salinity (Sundene, 1953). irradiance, transmittance through -the water and Moving southward along the west coast of Sweden, shading.!easurements of underwater light in the upper limit is progressively depressed the Isle of Man (Kein, 1966, 1971b) havc shown (Nichanek, private communication) to 15 ni or that in mid-winier the irradiance at the depth deeper.The plants are much smaller than usual of the deepest normal lower limit of L. hyper- (Kylin, 1907;Le-v-ring, 1953).The salinity at horca at 15 n below LAT is probably of -the same the inner limit of the species is probably FIRM/S87 Laminari_erborea between 25 and 3O6oS (Lcvring, 1953; ?'ichanek, was present) was saturated at about 300 rg. cal/ private communication).,In Loch Etive in cm2 sec in March and Nay.Ai the came time the Scotland, the inner limit of the species coin- light saturated gross photosynthetic rate cides with a salinity of about 26°/ooS (Powell, inreased in the new frond from about.8 ini 02/ unpublished). dm hour in March to around 3 ml 02/dinhour in Nay and August and was reduced to about Ir, the sites on the northeast coast of 1.3 ml 02/djn2 hour by November.In the old frond thgland studied by Bellanty e-t al. (in press) both the level waarothîd 1.8 ml 02/clrn2 hour in March the productivity arid the depth range of L. and May.The increase in suration level and jer,orea were found io be reduced in polluted gross photosynthesis in the summer were probably sites,In sites with turbid bu-t unpolluted water due to adaptation to increased irradiance in the the depth range was reduced less buthe produo- habitat.Lflning (1971) also calculated net -tivi-ty in shallow water was even lower,Reduo- photosynthesis at 100 and 400 rg., cal/cm2 sec a-t tianin productivity in both cases was attributed the same times of year.At the higher irradiance -to decreased light penetration,-The difference there was an increase in the new frond from March beiweên the groups was attributed either to the, to May from abc-u-t 1.5 to about 2.3 ml 02/dm2 hour more intermit-tent nature of the unpolluted tui'- arid this remained the same until August and then bidl.-ty, due to weather conditions, or to some dropped to about 1.2 ¡nl /du2 hour in November, relative stimulation of grow-th by nutrients in In the old frond the figures foMarch and Nay polluted water.The latter explanation seems were about 1,0 and 1.8 ml 0a/dm hour respec>- unlikely if light is limiting growth.There is tively,Thus, although respiration increases no evidence, in -thia area, of inhibition of the with highe' temperatures in the summer (see 2,42), growth of this species by polluting substances-- the high irradiance and adap-tation of the frond other than lhrough the reduction of light pene- allows plenty of assimilatory surplus a-t this tration.This is supported by experiments on -time of year,The lower figures for -the old the early, si-ages (Hopkin and Kainr 1971) in which frond are attribut-ed -to ontogene-tic change,At there was nc inhibition by water from the aver the lower irradiance, however, the crunmmer re- Tyrie estuary provided that the salinity was piration rates are relatively more important, so adjusted.Thrther laboratory experiments (Hopkin -the net photcynthesi.s falls from about and Kam, 1971) have shown that a number of 0,5 ml 02/dt hour in March to about half this in household detergents are not toxic at a level Nay and-August,Net photosynthesis of the old likely to be found in the seas though heavy metals frond at the low irradiarice is higher than the and a herbicide could be,After severe oil new frond because of lower respiration rates, pollution and subsequent detergent treatment on the southwest coast of Ehgland, no deleterious In a series of neat experiments, lAMing effect on populations of L -j*erborea could be (1969b, 1970a) has demonstrated how growth of detected (Bellanty et al., 197 4 -theew frond during the early months of the yedr can take place no-t only as a result of i-ta 2.4Thitri-tion arid growth own assimilation hirt using material from th id frond and stipe.He first showed (L«nmng, 1969b) 2,41Assjmjlaiion that the new fronds on plants on frames in the sea at 2.5 m developed the nmáximuxn area during It is assumed, in view of the lack of the fast growing season only if both old frond evidence to the contrary, that L. hyperborea is andstipe were present.,Plants with either of autotrophic, these amputated produced smaller new fronds., In whole plants kept in -the dark the now fronds The photosynthetic rate shown by zoosporas grew appreciably0He alec found that amputation or early gametophy-tes for the first two days of most of the frond in September affected the after release only just balances respiration size of the new -frond produced,1'rom -those even in saturating irradiarice (Kam, 1964).The observations he concluded that growth of the saturating irradiance for the growth of early new frond was partly at -the expense of material stages (see 2.3) is low compared with most from both the stipe and the old frond,This he plankionic end many benthic algae so far studied, took as an explanation for the observation that There have been no measurements of the rate of in young plants at least, each annual frond is photosynthesis (in terms of carbon fixed per normally larger than the last (see 2.44).In unit dry weight) of the early stages. later experiments (LUning, 1970a) on variously arnputa-ted plants in complete darkness new fronda LtTning (1971) measured the gross photo- grew only slightly when isolated from old froyids, synthesis of new arid old fronds at different with or without the stipe,With the old frond t:Lmes of year in relation to irrad.iarice.He present there was considerable growth, parti- found that the new frond was light saturated at cularly without the stipe being present.Thus about 400 »g. cal/cm2 sec in March at 4°C, at it seems that in darkness the ir-tipo does no-t about 500 »g. cal/cm2 sec in May at 8°C an supply storage material to the now frond but August at 16°C but at about 200 »g, cal/cmsec may parasitize it, for the production -of now in November at 10°C.The old frond (when a new one stipe and haptera,Lininig suggested that about FIR?4/S87 Laminaria Jyrborea 30 percent of the new frond area produced during show the high May rate, remainiag at about 0.2 rapid growth le due to storage material in the ml 02/g dry weight hour,High respiration was old frond.LUning (1971)iso showed that the -Lhus associated with fact growth and not with oid frond could contribute to the new frond temperature, suggesting adaptation to ambient growth through its own assimilation.He exposed sea temperature, the olçi frond t.labelled bicarbonate and light and found a gradation (from the collar downward) Rxperimen-ts on cell-free preparations of of activity in the new frond, whioh was isolated L. hyperborea (Johnston and Davios, 1969) have from contamination.In further experimente on indicated that many of the respiratory enzymes translocatlon, LUìiing et ai. (in press) placed present in other plantsan animals are present labelled bicarbonate in a email chamber glued io in -this alga,The actua.l biochemical pathways part of the illuminated frond,They found that could not yet be definedhowever, transport took place almost exclusively basi- petally and was confined to the large medullary 2,43Nutrition elements in a longitudinal strip corresponding to the position of the chamber.When the latter It is well known that species of Laminaria was on the old frond this active strip narrowed accumulate ions from sea water, some to quite at the collar and widened again in the new frond. high concentration factors, but there has been Transport was still active 24 hours after expo- little work on L. hyperborea. sure, but labelled material had mainly accumu- lated near the transition zone after 48 hours. Little is known of the nutritional require- The export of material outside the tissue exposed ments of this species,Ii la normally assumed to labelled bicarbonate (which was part of the that ni-trate and phosphate are necessary addi- old frond when this was present) did not exceed bions to sea water for laboratory experiments 1.5 peroentbut was much higher in April, during and there is little development of the early the fastest growth of the new frond, than in stages in etagoant sea water with no additions, January or October. Harnee (1932) added small quantities of potassium nitrate and phosphate to L, perborea The generai picture that emerges from these cultures every 10 days and investigated the experiments is as follows,At the start of the effect of different iodine additions.She fast growing season (see 3.31) in November or o].aimed that low concentrations of the latter December, the irradiance in the habitat may weil had a stimula-tory éffect,There has been no be below compensation (Llining, 1971;Kam, further evidence -that this species requires a 1971b) and growth takes place at the expense of higher concentration of iodine than is present storage material, mainly in the old frond.As in natural sea water,Growth of the early light conditions improve both old and new frond stages in the laboratory in Kam's (1964) medium assimilate and this material is incorporated of itural sea water enriched with 1,0 mN 1JO3, into the new frond,The highest assimilatory 0.1 mMK2HPO4, 0.005 eiN FeCi3 and eleven vitamins surplus is in the summer months (LUiiing, 1971) is as fast as has been recorded for Laminaria, and allows a larger frond than in the previous year in young plants with stipes which are self- 2,44Growth supporting in terms of assimilation,LUning (1970a) suggests that, because old stipes are Under optimum conditions in the laboratory heavily infested with epiphytes and epizoans, -the gainetophyles mature and produce gametes in assimilation may not be possible and these are about ten days from -the release of zoosporas. parasitic on the fronds,This could explain why In the case of the female, this may not inrolve older plants do not produce successively larger cell division but only differentiation and fronds each year, increase in size.The stales although smaller, always becomes multicellular,Under au's-optimal 2.42Respiration conditions maturation takes longer and there may be vegetative growth of -the gainetophytes prior LIIring (1971) measured respiration rates to gamete formation, of frommat different times of year.Taken on the hacia of rate of oxygen production per unit The first two divisions of the zygote occur of frond area, there was a slight increase from in quick succession but after -this cell divìeion March to August,However, as the frond thickness of the- sporophyte takes piace about every 45 changed during this time quite different results hours under optimum conditionri and while the were obtained on the baci n of dry weight,Then plant is still monoctromatic,This results in the value for the new frond increased from about a plant of 1 000 cells in 20 daya from spore 0,2 iii March to about 0,7 ml 02/g hour in Hay and release (Kam, 1969),Thxring this phase the dropped to about the Narch value in huguet and growth has been shown to be two-dimensional with December.Ac storage materials increase during slight increase in celi sise but very little the summer LUning also expressed these as volume change in plant chape (Kain1965),There is of oxygen per unit weight of crude protein.The therefore a straight-line relationship botbicen maximum was etui in May, being 6,9 ml O/g pro- the logarithme of sporophy'te length and cell tein hour respectively,The old frond did not number.The relationship is such -thai a 2:6 FII/S87 Laminaria hyperhorea sporophyte of 1 000 cells would be about 1 mm rate could be faster under optimum conditton, long (Kai.n, 1969).This species has not been possibly including weeding out of competitors. observed further in the laboratory but has been measured attached -to s1idein the sea (Kam, Further observations of' these cleared 1965).The plants were fdnd to continue blocks will be published at a later date.A few logarithmic growth in length at least up to measurements can be mentioned now.Table I lo mm.The vate of growth in length was about shows the stipe and frond lengths of tho largest 10 percent/day, half that observed tinder optiTmim plant of L. hyperborea removed from a 0,5 x 0.5 m conditions in the laboratory, but as the plant quadrat after the first, second and third year was by this -time mainly polystromatic, this rmst of growth on blocks cleared in November in three have involved a similar rate of cell division, successive years.'It also shows the lengths of A-t this observed rate of growth, it would take the largest of 20 plants remaining on iinsampled a spox-ophyte of 1 mm 25 days to reach 10 mm. parts of these blocks 'and measured in situ.The The total time from spore release would thus have growth rate after the first year was probably been 45 days.This rate of growth may rarely be nearer tO the maximum than during the first year achieved in the sea, because there was little competition from other species. There is less available information on the rate of growth from this stage onward,It is L1ning (1971) has followed the growth in difficult to find the máximum rate of cleared frond area and stipe length of plan-ta initially areas in the sea because of competition from nearly -two years old at a depth of 2,1ru below faster growing opportunists such as Saccorhisa LA!P for two'and a half years.Figure 5, taken and l)esrnare stia during -the first year. Kam directly from his paper, shows this,Increase (w;publishe(flas observed growth on cleared in a new f'ond area is apparent by Febi-iary and areas onth: submerged blocks of the roined there is a rapid increase in April and Nay which breakwater .t Port Erin, Isle of Nan.One block has ceased by July.During the slow growing was cleared in November in such a way that only season a considerable area is lost from the frond. plants of L:ss -then 10 mm could have remained, The frond produced in each subsequent year is From successive samplings from the block9 the largerthan the previous one,When LUning largest plant of L, hyperborea was measured, (1970a) grew second year plants on frames at The growth in length was roughly logarithmic for different depths, he found that the maximum frond about five months (after which competition was area was reached in June at 2 ru and. not until serious) and corresponded very approximately to July a-t 6 ru.The July frond area was the same 2 percent/day0As this was during the winter, at each depth but the stipes at 6 m were only a although in ehaliow'water (1.7 m below WiT), third of the length of those at2 in,The frond: light was probably limiting so the potential atipo ratio was therefore higher deeper.The growth rate must be faster,On another block situation was similar in third year plants, cleared in August, one plant showed a rate of at least 3 percent/day during the first three months. Jqhxi (1968) grew trae splatedfirst years' Thus9 at the fastest rate observed so far9 it (a-t -the s-tart) plants of L, perborea on frames would take a plant 145 days from snore release suspended in the sea at 1.3, 7.4 and 13.5 in below to reach 200 rum,There is no doubt that this LAT for a period of five months covering most of

TABLE I Stipe length + frond length in mm of largest plant in sample from cleared blocks at 2 ru below LAT at Port Erin, Isle of Meat

Block cleared, in November: 1 year 2 years

1967 270 + 400 340 + 760 1968 90 + .350 340 + 800

1969 100 + 410 Longest stipe length of pianti; measured in situ:

280 580 740 FIRM/SB? Lamjnariaerborea

HH25cm 25-Frond arca Frond 1969

20 E '5 Frond 1968

Frond 1967 Ì Frond - . . 1966 ... .. E _Sti'Iengh 20 o--oOc 10 s 1968 1969 J. F.MIA M!J.J. A S. O. N D J. F,MIA.MIJ. J. A. S.O.N,DJIF

Fig. 5Records of the growth of frond and stipe of L, hyperborea at 2 n below LAT iñ Helgoland (from llning, 1971). the fast. growing period0As the old fronds were the stipe and frond weights and ageing of the cut off before the experiment, growth may nat have stipe base (see 1.32),This method has been used been maximal. At 1.3 n by Bellamr et al, (in presa) in terms of dry the stipe length about doubled, at 7.4 n it weight and by Kam(1971a) ih terms of fresh increased by about half and at 13,5 n there was weight. n increment is calculated for each year almost no atipe growth,The frond, however, grew of life, remembering that the stipe is perennial at all depths, at 1.3 m to nearly 10 times its and the frond annual.Thus the s-bipe increment original length, at 7,4 n to about five times is taken as the difference between the mean stije and at 13.5 n to about 2,5 times,These figurEs weight of an age group and that of the previous are very approximate because of individual age group and the frond increment as the mean variation but the differences between the depths frond weight for that age group.The recul-ta are clear,The results agree with Ltlning's in for each successive age group are then added that there is more reduction in stipe than frond together.The population should, of couraebe growth with increasing depth. sampled at the end of the fast growing soasan when fresh weight is used bu-t later for dry Growth of the frond of this species can be weight.Some cumulative fresh weights for followed by observation of holes punohed In it various populations calculated by this method at various distances from the transition zones as are shown in Figuro 6.The. di sadvantageof has been done in other members of the Laminariales the method are first that the error9 toe9 is (Parke, 1948;Sundene, 1964;Norton and Burrows, cumulative and second thai any one year may not 1969). be repreaentative, The past growth of plants of a population can be assessed ai any one time by measurementof 4000

1000

500

100

¡ - I j J 3 4 5 6 7 8 9 10 Age (years)

Pig. 6Cumulative fresh wei.it production in populations in the laie of Nan andthEspegrend area (see Tables IX and IIfor positionsand depths).

Q,s4 A,BM ¿,ME4 ., D ,)1E14 BE33 ME22 BCuO.5 , , ( 'a Kam, 1971a) FIE4JS87 Laminaria h,yperhorea

3 LIFE HISTORY conditions in the laboratory there are at least five paie cells by the time the gametophy-te is 3.1Life cycle mature, within 10 days from spore liberation (Fig. 7, left).Any coil can become an anther-i- 3.11Alternatiòn of generations dium, the apex of which swells and fortes a cap, bursts and releases a single antherozoid from L. h'perborea shows marked ho-t eromorphio the gametangium (Williams, 1921).Schreiber al-bei'nation of generations typical of the order. (1930) found. that maie survival was favoured over The large habitat dominating sporophy-te alter'- female in prolonged cultures, perhaps affected by ns-tea with microscopio dioecious gametophy-tea increasing salinity with evaporation. which have not yet been observed in the field. libr a while -the effective coil of the fe 3.12Nuolear phases gametophy-te merely swells spherically.It may then divide but under optimum conditions usuaJ.ly Evans (1965) showed -that the gametophy-tes does not,It then elongates, to -the shape shown were haploid. with aboirt 31 chromosomes and young in Figure 7 (right).The tip of the process sporophy-tea, providing there had been fertili- thickens and chromatophores crowd. into it za-bion, diloid with about 62 chromosomes. (Williams, '1921).A slit then appears in the tip Schreiber (1930), after separating gainetophytes, and. -the contents of the oogoniunl paas ou-t through showed that unfertilized eggs could develop into the gaping opening.The egg is spherical end parthenogenetic eporophy-tes which survive for a about 12 p in diameter. The end wall of -the now limited period,This ham been confirmed empty oogonium closes and forms a cup-shaped p1a- (Svendaen nd Kam, 1971).Evans (1965) had form on which the egg may rest.Williams (1921) one countf 36 chromosomes in a young spore-. was the first -to confirm fertilization though he phy-teof this species.This could represent did no-t actually see i-t.He saw eggs surrounded the haploid number and be explained by parUieno- by active an-therozoids and nuclei in various gene-tic development of that sporophy-be. phases of fusion,After fertilization by a male gamete a wail forms round thergote.The con- 32 Reproduction tinued development of the sporophyte is shown in Figure 8. The zoospore is about 5p in diameter (Sauvagean, 1913), arid is pear shaped with two Although zygotes are formed within 10 days lateral flagella arid. a curved chromatophore in -the laboratory under optimum conditions, (Williams, 1921),Sauvagean (1918) found no eye- etophy-tes cari develop quite differently under spot though he did. find one in Saccorhiza.Kanda other conditions,Schreiber (1930) maintained (1936, 1944) found none in severa]. species of his cultures in sea water with only nitrate and: Laminaria.,However, Kuckuck (1912) showed an phosphate added in en unheated room in reduced eyeapot in L.accharina zoospores.As Saccar- daylight,His cultures were unialgal arid devoid hiza zoospores are photo-tactic and those of north of .,rotozoa;this he achieved by careful washing hropean species of Laminaria are not (Kam, of individual early gaxaetopliytea.Hua gameto- 1969), it seems likely that Sauvageau was right. phy-tes developed extensively vegee.-bively, each one eventually becoming a -tangled mass of fila- Liberated zoospores can remain motile for mento (broader in -the female tbei -bha maie) up to a day in -the labors-tory and when motility several millimetres acreas arid thus macroscopic. is lost they round off, form a oeil wall and may Kein (1964) found that similar filamp.ntous remain planlctonic for some days, slowly sinking gainetophytes developed in unchanged medium (ICain,1964).They are fairly sticky and adhere providing that the irradieiice was low.Figure 9 -to surfaces,After rounding off, the cell shows these at an early stage in development.In extrudes a germination tube (Fig. 7) in-to which saturating irradiance the gamotophy-tes become nos-t of the cytoplasm and the chromatophore, mature before whatever factor that may cause divided into two, pass (Williams, 1921).The filainentous development :Ln unchanged medium tube swells a-b -the end to at least the size of becomes effective,Filaiaentous gametophytes of the original spore, the nucleus divides and one Schreiber' s type have been maintained for a few passes into the enlargement with most of the years in low light at 1000 (Svendaen end. Kein, cytoplasm arid chronatophores arid a wall forms 1971).The conditions causing this vegetative across the tube (Williams, 1921).The nucleus growth have yet to be defined.. arid cytoplasm remaining in the original spore degenerate.This is called the dumb-bell stage Filainentous gametophy-tes may be entirely and it can be achieved in the dark and while vegetative, may have occasional gainotangia or s-till plank-tonic. under the right conditions may produce many gametangia.There is -thus a far greater poteri- From -this stage onward the development of -tial in terms of gametes in this type of gameto- the malo and the female gametophy-te differs.In phyte,Under optimum laboratory conditions a the male the enlargement swells but soon divides female produces only one egg and thon dies, whi1 into several small cells ii-i a cluster (Fig. 7, filainentous gzunetophy-tee con produce thoucanda left).These divide further and under optimum over a period of years.The latter cari also be FIRI4 87 Laminariaj.erbo ea o 5OH D O flg

.g. 7Drawin from culture under optimum conditionu of development from zooaporea (top) of male (left) and female (right) etophytea to maturitythe production of a zrgote (third row) and ita initial development an a eporophte (fourth row). ?IR? SBLaminaria erbo rea

02mm

2mm

Fig.8Drawin of developing uporophytea.Top row from Savageau (1918). m - monootromatia p - polyErtrotic FIILj Laniinar±a hyperborea fragmented and the various parts continue the explanation might also be that minute sporo- growth arid reproduction. phy-tea were lying dormant until able to grow in increased irradiance (due to the clearing). The factors determining gamete formation in filamentous gametophy-teu remain obscure.In The spherical zygote is considerably denser noverai experiments Schreiber (1930) found that than sea water and if' it is not stuck to 'the gametes vere formed in winter or in artificially oogoniuin neck it could fail away from a. very induced low temperatures of 2 to 400 but were steep or overhanging surface.This may be why sterile at higher summer temperatures.However, the species is absent from overhangs, though the in saturating irradiamos,ametes are freely behaviour of the zoosporas could be responsible. formed a-t 18°C (Kam, 1969) and f'ilaznen-tous If the zygote is in the proximity of the oogoniwn gnmetophrten become rapidly fertile at 10°C In in culture when i-t produces Its first rhizoid new medium in saturating irradiaiice (Svendsen this usually attaches' on 'to the neck (Sauvageau, end Kam, 1911).Thus, there seems to be en 1918). Interaction of factors de-terinining the onnet of fertility in L. hyperborea gametoph,y-tes. The development of 'the sporopliyte from the zygote has been followed in -the early stages by It in not knowa what type of garnetophyte Sauvagean (1918) and others.The zygote elon- exists in the field and how in ni-tn conditions gates and the cross wail divides it into an affect the morphology.'Sterlized stones left apical and a banal col]. (Fig, 7, bottom row), at 2 m below MT on Port Erin breakwater for The next division cari be in either direction, -two months at different times of year and then resulting in four celle in a row or in a oruoiate examined uider a microscope bore L. hyperborea form.For some time cell divisions 'tend to be sporelinge only during the aporiag season, as simultaneous so that many groupa of four or 16 might be 'xpeoted (Kain, unpublished),On the cells can be soon to be aggregated (Fig. 8).At adjacent ilock, however, which was scraped eler,,r this stage the plant remains monontroniatio.At -two month, sporelings of -this species arose all approximately the 30-oeil stage a rhizoid is through the year.This suggests that gameto- formed from a basal cell arid more develop later, phytes might have been present all the time but The shape of 'the plant is roughly elliptical

500 ru

Fig. 9Drawings of filwncntoiw malo (left) and female (centre and, right) gwnotophrton grown in culture in suboptimai light without change of medium, FII/$8ImjnarIa Irborect (Fig. 8).As development procoeds, -there is The sporing season has been recorded in periclinal as well as anticlinal division at the Isle of Man by Kam(unpublished).The the base of the plant, resulting in a polystro- earliest fertile fronds appear in tizo middle of ma-tic region which will become the stipe. September and the latest disappear near the Gradually the base of -the frond also becomes middle of April.Between these dMes -there is polystromatic (Fig. 9 p) end the stipe terete. a steady increase to about 40 percent of mature At some s-tage (at about -two months in L. plants sporing a-t the beginning of January and ta according to Killian, 1911) growth then a steady declino.Labelled plants observed becomes localized mainly in the transition zone at intervals spored for a mean of 6.5 weeks each though also in the surface layers as more of but there was considerable variation in -the the frond becomes polystromatic, being -two, three duration.Ou-t of 160 only four plants spored and four layered and leaving only a tip of the over two separate periods and in each case the original monos-tromatic tissue.A diEc is formed second sorus to appear was small and, on part of at the base of the stipe and. this je fixed to -the frond that had. not previously borne -the substratum by rhizoids on its lower surface. aporaxigza.Parke (unpublished) recorded the Eventually medulla cells are formed from coz'- sporing of labelled plants near low water in tical cello in the stipe and frond. in the way Devon, southwest Jthgland and found a later and already described (see 1,32).Later, the first more in-tense peak to the sporing season in whorl of small haptera grow out above tIzo disc D'ebruary.Roes (1928) also recorded a later and. form further attachment. season, of February -to May with a peak at the

end. of Yiarch, in 'dales,The Isle of Idian obser-- Whether or not growth slows during the vations have been made over a period of 12 years first slow growing season depends on when a young and differ only slightly from year to year so sporophy-teroso and probably on conditions. the differences between these sots of observations must be due to the locality or, more It ¿an be seen from the developmental likely, to the selection of populations in stages in tzo figatros that the frond:stipe ratio Devon and Wales..Even in the Isle of Man it is high in rery young sporophtes, even before baa been observed. thais a group of plants of the shedding of the first frond, very s.milar ago may show a closer timing in their fertile periods than the general popula' In normal growth the sporophyte of L. 'tion, and -the peak may be in a cliffcrezzt month. y-perborea obligatorily has only one holdfast, It appears that near the north of u.s rango too stipe and frond.New stipes cannoi arise from this species produces spores inixztor (Kam, the holdfat as they can in some species in the 197 la). genus.There is thus no vegetative reproduotion of this phase.The region of paramount inapor- Nothing is known of the time of day of tance is the transition zone.If this geta spore emission. damaged in a particular way, a freak frond can develop, and if it is split longitudinally into Sari develop only on the old frond and in two without too much other damage, the divided mature plants cover all but the distal and a-tipe and frond can continue growth in two parts. proximal paz-to (fig. lOA),The sorcts on each If -the zone is cut transversely, losing the finger of a digitate frond extends to about 2 frond, the stipe can continue gro'sth for a short from the lateral edge but there may be 50 mm or while only (Kam, unpublished), during the more of sterile frond a-t the tip.Near the growing season.If -the transition zone is transition zone, or after the growth of' the new destroyed, then the plant is incapable of fur- frond, the collar between the fronds, there is ther grow-tb and dies. an arc of s-tertio tissue,There is a sorus on

each surface of the frond and these are nor-- The earliest age at which a sporophyte of inally exactly opposite each other.The coverage L. hyperborea has been observed to become fer- of mature frond surfaces devoid of epiphytos is tile is about 15 months (Kam, unpublished) but 73 percent (Kam, unpublished).In young fronda this is unusual.Some have been observed ai 18 sporing for the first -time, the sari are smaller unths and almost all plants on cleared areas at in relation to the frond and there is more of a 2 a below TA'T' which were two years old near the p on -the proximal part,I-b is here that a beginning of the sporing seanon have been second fertile .,rea occasionally appears after observed -to spore.However, the maturity of the -the first son have disappeared. aporopbyte probably depends on size as well as age, so in a forest where growth is slower than Certain epizoide and epìpbytea affect the on cleared areas, fertility may well be arrived development of fertile tissue.Sari, have boon at later.This is supported by.thobservation. observed extending only up to about 2 nun froto that two-year olds on cleared areas in deeper the active edge of Membranipora.The other øidO water (6 m), where growth is slower, did not of the frond can apparently be affected by active become fertile during -the sporing season.0f grow-th of this bryozoan.Sarna formation can

100 marked plants in virgin forest only 75 per-- also be inhibited by small epiphy-tos. However, cent of the four and flve.-yeaa' elda spored while epiphytea can be found. growing ou the - curface of all those of six and over did (Kam, unpublished). a mature sorus (Jaasund, 1965). A

Sortis

Parophyses

5Opm Sporangia

Fig. 10The aorte in L. hyperboroa A. Dratinof fortilo frond.Stippled B. Pranaveroe oection of aoriie ohowing aroQg .ri on bothurfaeea. uniloonlar aporengia end paraphyies. FIL87 Lanin aria h,yperboro :7

The eoruswbf,ch develo fta th? 3UC October (Svencthen end Kein,1971).'However, ficial oeils of the :rood.i: ri :ue of club most of the old fronds are oeualiy lostin March shaped paraphyoc; u1( ii c., 'to Na ...... one ai-to those in shallow ter (Fig, loB) rJrJ,. .yrr0r l''rc alat ocni$ go hefoi:...deeper once (LUning1970e).'The thickenthgs at their dr 'ah :ìdhero az'rjvai onhe bench of a sacs of fronds is together and. form o. ioug; r> lohe often culled, the "lAny cact".' sporacg-1a0The latter aim oidmrora]. chapad andan usually he coco in .lurcj a-Laceo of t' j m'i Idi frond grorrththe ottpo developncnt0the1;her there arerogreecirc elongutee3--:,uI' a'Icìt't'cm bIc clj,ebmmd.e-ecl by crops of the as dialmeci foi' L0 obarc. the Iraucli tOO w, eti:, is pio ud (Parke1948) ii no-t There a'o obow; snootAm2 it bitag'oumcsomilbc' hoí'mi'.r 'the papillon 7 300 opommgia (inoiuling c.0relopiri. oner4 per on itS surface oppar 'cud Ghoce 'too are paler square írdliìotetre of frond 'face area0 then -tAie oldcoon.' 'Ihm' or' part of the s'tipe ta Schreiber (1930) found. that othe 32 zoospores devoid of ep:Lçhytao ?oi eo;ie 'tise, produced in each eporengiuinhalfwerepotential male and half pa-ten-tb). femo1 ogr'rii-oprtcc0 Thenew growth of 1amcpteraJim a ring above Thus seis genohypicntl c-hreibao 'thelast. 1mo i p,e rarI bl,e in i iotiming.' 'The described aporangial Cte3.o'oco0Thu con-bents fire-t knobs ¡soy ctm'o' s' in Jnmeary and, the lìírp-.' surrounded by a membrane9 cooll; with a jerk from iba T'rI' Lo;i b' rol'o" -they mo,',r not the sporndum cad moveolowlyoutward 'through apuea2' until ,TnJT,The railler appearance may the paraphyeesp:-3h>;:r.: ...... ;chcd. by thetr prec be confined 'lo trorwama e!an'tc end soma il to:' sure, leaving ithe emp- era-ogiwn utili attached0 plante do rio-t praduce lwp'iera every mear.' The ball of cooaporeu::omneins thtact outuid,e 'the paraphysea for some seconds and then arm opening I;ho shedding of 'the old fiwmdthe appears. cab 'the 000sporec sria out0 p:t: ' ir ru3.]y f:mi.rly free of enipby'tes and epicoi. : from the tz-nmiui'tian zone upward for a 3.' 3 while0The Lambeta forest :.'r June end July thne, has a rel,a'tivo]y tm'oieoa" m.oPearance, with 33i S';...... :.::ia;ion in anternal mmct of the smooth brown,.:. o:r- rooluttared by e ml mo::phology other organicen apart fz'om or.eilucida. The highly smneilagi,ioowms ,i:':e. surface is not a Because io t 3 r' c oj'eo-rao.:'J. ert ve_i,( bosmoi-t'bio r_ol,rt 'o,tvs "omsnat oo'gc'n'l curo the seasonal oheuger, o-.'oi;iyo .ur1idrai °m'ecemeati,ïh-sss'mvor9(sroii'el i:1m ?con-la c'ra iporpbo'i ogy' ra',he'thea' ir pci'm' m 0.4, on :1.-ito» aotc'oi:cd. hg," b licJ, ryarmrrs hommCr,? tporzi Thece cbarßoa ai-e aaOcim9ri icih IhowC.*CCt r',emmb-cnoenac ,:id ,1, cm c'tec'''I,'I-'it iiu r'e-feco of varia'tfou in grow-th i>-,',c w")r uerr,,(n.Tlo thzt:3I a nace ,î"-' o Ciii ,:-çh't'20, r' tire soc tt-ler,ieul; growing sanean sta-to in Toi mmtho t of o'pJ. cIr; a,'-,, T';:' hc' an:(mí' i ho 35,sr'20r9»arti'' 'transition roue bececiec paluraid o. C- 'ter) -' r:r'l J,cr st'i"m. S'rc',O s'the heeds swelling acpae.rc 4j) b.3e of 'the frond0The ear- a oaims, ç1a-ll) 1,2 Ji> 11.:,t1; the '7l'2'Iez0 onset is remarkably t iJaoe a-r-thg 01'')lÀ"3'ai cf the a,-, '.c1r'I:,ar'0i19 sono within week's of caz-ì ojccm'0trT:,rj thlb. r.', tl'a 2'.o' 'cr I 'o, -, 's1: a:':',1i 'rie and is the new frondm ml'mw3lmr :D9d;:uto 00u'jlkJ:i 0001e ¡5 ?icir,'1r'-- amm .75)30i'ottie' cuJ.comico.ìion the shape of a 'tùiamba9m'on 'ith,brmatlyc oy ooiplts"hoI.07L2 1J,'.'I3 »17 c'siihr; pe:Hc; that below and the old L'i-,.nd obmne0 hic.r3o kerl haca fetle, TaCrmra 'ornocl -trIoni. ne,cion with the oid frond is narorwer -than the tiro,eurio.;. er'a,ic,IJjii't',-bs,'J. new frond the correbetweenthey 'ic'a ho called :1,32 £1OC1'. '0d.,117) ir0jh, -thecollar.' 11e :>::;ce i ìm..:...'-..',er,to'tic cf -the species gz,rmm r.-' rtd.o.:1 d.- uii->g candi' The te'mal frech micigh't ci' any rnn:l,vro pleut 'tiens.'The mew frrnd e:;ïu'od thcr (C'.',ig, IB) varie.. :nimuiderai'iy wb'th -the osaron beecuso o' end becomes prograu,ral; 'hu. ï'' mmsme..ina ti'...... rol, developwwi'i of ' the froci'i.'The mant.'.' paler -than the oid. ftc'md.r . ['rc'h rodac' mum r;be1otained rasivi tuna rlarrrig the arcing9 turbulent eondi-t:L0009'c >'l cpi Io haire juS-i 'fore -the beer cf the e1f.rnd or nac ho appeared tu the f:3n? 30r 'o çir. or ffîgorri say in. Jt.y ctheo -lut o-rrr'u-mP te ihiJy grown.'The havebeen .'rced O' Jc' rJ'' (T3g0IB), By minisawi sony he r> IL-r re,nbC-'2 after come losses-to flprii 'theoe'l' und, ompmaio'r --h-?ciJ cc' of the tho foowi b,' ',e'wirs a -tmrn,og ocmi Juai, be,COiO old,By July grow-b br>' Oe.'s(t Thb2 tAie go:to;b o rtira ncr ocmi or io U'e opring jiii, of events tuk'oo'tc',ce .'mnc;-th'.r.Lec 5r deeiormr - afiar 1ko baca '-n''iba aid, :'rond,b" ejjsmuns iIj water timer. abmrllcu (i Tn i9'(U.-',). TAie 'time iseigh'i j-e Dz2 tei:is f:er'ispm,eiihor to tiovembox' of separectioa o,? 'the,ld. ír,'md. fran tAie oar' (Ilbi i,ckm 9.S9) td.'ro inc -C'rnod con-laine a depende o» wc-ter rrmrnc,ut 'L'l,lo'T lonenda "n comcplerimr'i o" c'io ,ro ww'6owia 'I1ù tIJ ch fourcl -the degrmo of sha!irao" o '.a.e'ii.m;ro. '.hr dopth thathe rmy ;aj '-Iat' im tact f:cedi_' in deep end -the incidence ci;tom;o.' tear, tIn's vz"y tenter foi L ?oatoa. 'the :m'r tarbon tirano in considerably frasi year 'to year at orme cite0 'aballct; cra'icro.'.re,aor,aai,I:r boiorea ai' t'cractef 'noc Under turbulent cowtitiomas the eid içnnrt can be of oa,-rczrn-,o mr tar c'L (o 2,i) Aicelaenle-a! J,or" lor.'t tu curly I&-rch ;:blo -ar'- o1roci-n1 olmi oh 2co'a'iOa eoa'°r- :.ocha ?s''tCc' Arr churl ltot francio of r,cumitia,o c,c'r' ho '.-ecatrmod until water, FI1 S8 Laminar a h a erborea

4 POFILJLATION populations, MCa4 end BE33, wore the stipes 4.1Structure all significantly smaller than in other popula- tiens.In both this may have been because of 4.11Age compòition reduced light, the former being atimn below LAT bui at the inner limit of the species in a The age composition of a population cave, and the latter being at 33 s near the obviously deponds on the history of the popula-. depth limit.However, in the former cases the tien and the environmental factors. ypes of type of water movement at this site, which population can be broadly grouped in this res- caused. damage to the mneristem, may have been pect.Figure 11 shows the age structure of responsible for the small size,Small plants some of the populations studied by iain (1963 in deep water have been reported by Prints end unpublished),In the top lino are populations (1926) in Trondheimnfjord.The stipe end frond from shallow water exposed to considerable wave weights of the plants in the Isle of 1an sites action,They all shöw a predominance of young from 1 to 6 m below LAT d not differ greatly. plants, presumably made possible by the continual Differences in one-year olds may only reflect removal of larger plants by wave action.The temporary conditions or the time of arising of similarity of F2, collected samples from &if- most of the group and as size distribution is feront sites in the Isle of Nan at 2 mn below unlikely to be normal, fiducial limits cannot LAP made in 1959, and 1tE4, collected from a single be relied upon.The lower stipe weight of the site at i mn in 1966, encourages a generalization slightly more sheltered MS4, compared. with of this sort.In the second line are more MF4, may possibly be explained by the frond: sheltered siallow water populations.Here one stipe ratio being increased in sheltered. water, would expect that without regular removal of This is shown much more clearly in B54 (L. large p1ans, more intermittent factors might hyperborea f. cucullata) in which the stipea allow the tevelopment of younger plants.Thus were very much smaller and the fronds larger various agn dominate.In the third line are than òther populations,The fact that down to deeper populations where conditions allow thick 20mnor so in these populations there is litt]o forests to form.This is the most stable situa- x'eduòtion in stipe and frond weight is probably tion because storm damage is minimized and a explained by the interaction of density and broad age range made possible.This is shown depth (see 4.22),The largest plants for their well in 5H10 and in BH14 but not in F5.The age in these populations were those of BCuO.5, lastwas perhaps shallow enough fox' the moi'- in a fast current near Bergen.It is likely tality io be affected by wave action.Finally, that by some accident the site had been cleared in the bottom lineare populations from the of vegetation some years previously, because edge of the forest, not mach affected by the the plants were all young, and that the environ- canopy, where the critical factor is onò affeo- ment was favourable for fast growth.Recent ting the establishment of the species on the obiervations (Kam, unpublished) on cleared rock,Here intermittent factors are all- areas have indicated much faster growth rates important and establishment is severely affected than in established forests, by the conditions in a particular year,In both 1962 (L2 and Lb) and 1966 (ME14) there Tai,lee IV and V show dry weights found seemed to have been a good establishment year by Whittick (1969) of atipés and fronds res- four years previously. pectively, using the sano method of aging as Kam, at four sites in Britain.It is clear 4.12WeIght or size composition from Tables II to V that there is more variation in size within an area and thus with The method of aging plants of L. hyperborea local conditions than there is betwen the instigated by Parke (unpublished) allows a widely differing latitudes observed. determination of the size/age relationship within populations.For some time it was thought This is also apparent from Grenager' s (Kam, 1963) that the stipe, being perennial, is (1953, 1954, 1955, 1956, 1964) data.His a good measure of the growth of the plant through- measurements were all on plants of L. hyper- oui its life,However, it was later found. (Kamm, borea which were brought up whole by the spring 1971a) that environmental factors can have a grab he used for surveys of quite large areas marked effect on the frond:stipe ratio so miei'- in Norway.The surveys were made after the pretations must be made with care when certain end of the fast growing season,The plants conditions differ,Table II shows the mean stipe came from a wide range of depths enditein fresh wetghte (with haptera removed) with fida- each cases so each sot of observations is more ciallimnits (p0,05) for different ages in collective then those by Hain,He counted the widely differing populations, taken from Kam number of "rings" in the stipe.This is noi (1963, 1967 and unpublished).Similarly Table quite as reliable an estimate of ago as a lon- III shows the frond. fresh weights.Some of the gitudinal section but may often give the same data published in 1967 involved an overestimate result.lUs data for the mean total frech of the plant age (Kn, 1971a). In only two weight and total length in plants with different number of rings in a number of areas 4:2 -.

ME4

1112 30 MS4 sS BS4 2

F5 SF10 B H 14

1i2l3

LlO M E 14

I 2 3 4 5 6 7 8 9 10 2 3 4 5 6 7Ï Age in years

p9g. 11 Age structure of populations, Percentages of plants within agegroups, See Palles IX and II for ai-Le positions and depths, (From Kam, unimblished.) YI8M/S81Laminaria hperborea

TELEn Mewt fresh weights of stipes at various ages in different populations sampled by Kam (1963, 1971a and unpublished).I'iduoial limits at 0.05 shown in brackets. For latitudes and longitudes see Pable IX,

Depth Minimum in years Sei m below LAP 1 2 3 4 5 6 7

MCa4 1 0,697 5,32 6.47 6.44 14.7 (±0.383) (±2.07) (±2.43) (±2.99)(±1.67)(±10.4) MF4 1.89 17.4 45.3 125 236 355 424 (65) (±°is) (±6.70) (±10.8) (±57) (±112) (57) MS4 1 0.906 7.34 45.8 119 132 224 298 313 348 (±0.467)(±2.60)(±12.1) (+14) (25) (±45) (4)(i) F2 2 22.0 52.2 136 gii 425 521 553 544 (12.2) (±14.7) (±48) (±61) (±75) (±56) (+76) (+81) F5 5 9,62 30.9 61.0 22'7 722 (±3.67) (±11,6) (±18,0) (±104) (±203) L5 5 26,9 78.8 223 313 319 378 419 (±5.8) (±25.4) (+67) (±46) (+67) (+6o) (+110) LlO 10 21.2 35.7 143 (±4.5) (±s.4) lEI4 11 0.604 2.41 27,8 79.3 110 208 (+0.426)(+0.83)(+12,3) (±0.8) (27) (±7) 11E22 19 2.61 7.42 13.8 (±093) (±3.71) (+8.) 3S5 3 33.8 46.9 86.6 225 (±11.0)(±'33)(±17.2) (±69) 3 181 263V 247 (53) (±116) SE16 14 120 (±52) BCUO.5 O 38.7 92.3 192 (±12.5)(32.7) BE4 3.5 0,290 3.60 529 733 (±0.091)(±1.57) (±153) (±105) BS4 3,5 4.91 24,2 32.3 61.7 71 (±3.89) (±6.4) (±7' 7) (±36.1) (63) BR14 14 0.821 5.10 27.9 73,3 115 760 949 (±1.44) (±1.58) (±13,5)(±23.6) (o) (47) (+i) 18 18 30 61.4 87.1 (+13) (±18.4)(±44.2)

flE33 33 4.46 4.56 (±2.62)(±3.87) T2 i 4.61 18.2 (±3.22) (6.2) 4:4 FIi1/S87 Laminaria hyperborea

TABLE III

Meanfresh weights of fronds at various ages in different populations sampled by Kam(1971 and unpublished).Fiducial limits at 0.05 shown in brackets. For latitudes end longitudes seo Table IX

Minimum age in years Site Depth m below LAP i 2 3 4 5 6 7 8

MJf4 7.23 43.9 81.8 181 221 288 320 (±2.39)(±15.1) (±28.1) (±74) (83) (76) (±132) 5.09 25.5 67.3 203 287 277 421 (±2.3 1) (±73)(±34.2) (4o) (±121) (±2°) (±159) ME14 11 4,74 10.3 55.2 108 120 258 (±2,17) (5.1) (±21.5) (15) (±42) (±111) ME22 19 15.1 29.3 37.4 45.9 (5.o) (±10,9) (±19.6) (±13.4) BOuO.5 o 132 279 616 (62) (±1«) (±187) BE4. 3,5 2.32 12.0 54.6 133 546 796 (±0.55) (±6.8) (±31.8) (±24)(244) (±194) BS4 3.5 87.7 159 182 288 (±64.2) (ï.i) (+61) (9i) B1114 14 10.4 33,4 54.1 92.0 119 431 605 (±2.8) (±10.2) (±15.2) (±22.3) (32) (84)(54) BS18 18 182 344 274 (80) (±156) (±108). BE33 33 14.2 14.8 (8.8) (±9.8)

TE2 1 16.7 99.0 (±11.8) (8.5) are shown replot-tod in Figure 12.It seems thai 150 cm,This difference could have been due to the fresh wei.*t approximates to a straight -the Scottish sample being from a small (favour'-. line rather than the more usual siginoid curve able) depth range, to particularly favourable for plant size against age.This may be due to conditions at the site, to a difference in the the mixed nature of the samples,Plants in deep method of counting rings or to the relatively water grow more slowly and may riot live as long small sample size. as those at an optimumdepth, Theirmean fresh wei..t will thus reduce -the mean for plants of The maxiirrum length attained varies to sosie medium age but no-t the older, largest ones. A extent with latitude. Sauvagenu (1918) stated similar effect may be produced by a dense canopy. that ma.zimswnstipe length in Francewas usually Although Grenager'ssurveys extended from south about I mnand the frond lengthrarely exceeded Norway to near the northeastern limit on the i a.This is true also of the Isle of )an; mainland, there seemed no trend with latitude few stipes have been found measuring more than of either total lengthor freshweight, at given imn (Kam, 1963, 1971).However, in Scotland age. s-tipes of up to 2 m (Kamm, unpubliehed) or even 3 m (Parke, unpublished) have beenrecorded. Black et al,(1959) collected asampleof Grenager measured plants fromhis sampling 112 plants of L. hyperhorea fromabout 24 km55W sites (see Table IX)and the longest stipe of Oben, western Scotland, measured the stipes lengths, with whole plantlengthin brackets, and oboerved rings,The mean stipe lengthswith were as follows: KvitØy, 1.35 m (2.0);Karmiyy, 2.2 m3.1 2.5 m (3.0); Iord-1'r4ya, 7 to 10 rings ranged from174 to 192 cm, whereas ; Tustna, -the mean stipe lengths with7 to 10 rings in 2.0 m3.2 S. Helgoland, 2.3 m (3.3); Hci,y, five of Grenager's surveys ranged from 78 to 1.3 m2.1 ; E. Finmark, 1.5m (2.4). FIRM/387 Laminaria h,yperborea

TABLE IV

Dry weights of stipes at various a«esindifferent populations sampled by Whittick(1969). Forlatitudesandlongitudes see Table IX0

Depth Agein years Sito m below LItT 1 2 4 5 9

Seimen 2 8.5 22 45 46 Cove (4.3) (+6) (+6) (4) 6 2.4 11' 19 38 51 76 (+1.2) (±3) (±4) (+2) (±3) (+14) 9 3.1 12 22 32 39 45 (±1.4) (±2) (±2) (±10) (±14) (±13) 12 17.4 29 33 34 (s) (±3) (ii) (-i-6) Dunnanus i 0.51 0.87 4 20.8 42.7 59.6 60.3 Bay (+0.18) +0.68 (±2) (±11.8)(±12.8) (±15) (±0.2) 4 0.4 2.8 10.6 27 57 67 (±0.i) (±°4) (±53) (±2) (±15) (v.6) 11 21 25 49 (±) (6) (+10) 17 0.56 1.6 4.8 6.4 18 (±0.24)(±0.6) (+1.4) (+1.3) () ìambo 0.7 5.1 5.7 16.4 22 25.6 (±0,6) (±1.3) (±6.0) (±3) (±4.2) 4 3 7. 42,3

(±1) (±2) . (±7.6) 7 4.2 9.0 16.4 26,6 (±2) (±2.9) (±6.3) (±7,8)

Petticoo o 0.5 2.0 8.9 21 45. 61 64 Wick (+0.2) (±°4)(2.6)(.8) (5) (+6) (6) Bay i 0.25 1.9 7,5 31.7 59 71.6 89,6 101 105 (+0.2) (-0.8) (±2.8) (±4.6) (±4) (±4,2) (±6.0)(f18)(±4°) 5 0.125 0.35 2.8 14.9 77 96.3 111 114 (±°°3)(+0.12)(±°53)(±45) (±10) (±10.3) (±11)(2o) 9 0.022 0.62 8.8 22.9 41 60 71 (±°°3) (±03)(2.2) (±2.8) (±4) (±7) (6) il 0.87 2,2 4.6 6.9 6.8 11.3 (±0.42)(±043) (±°°4'(±'') (±1.4) (±0.8) 4:6 F1/S87 Laminaria hyperborca

TLBLE V Dry weights offrondsat the peak season at variou.e ages in different pc'pulationa sampled by hittick (1969).Fiducial limits at 0.05 shown in brackets, Fbi' latitudes and longitudes see Table IX.

Depth fige in years Sit e n below i 2 3 4 5

Seniien 2 16.5 37 98 103 107 Cove (+2.5) (±10) (±48) (+31) (±11) 6 22 82 100 142 (±4) (±") (±) (+) 9 5.6 12.5 23 56 92 117 (±°7) (±2.5) (i) (+io) (±1?) (±15) 12 7.5 17.6 27.5 42 57 (±3.2) (±5) (±5) (±17) (±17) Duninanjw 1,7 85 122 155 Bay (±0.6) (+19) (+io) (,5) 4 2,2 28 103 (±10) (±12) 11 33.3 76 90 (±3.3) (+11) (+13) 11 0.8 1,2 42 10 17.2 (+0.8) (±4) (±8) (+1.8) Flamborou 0.1 lT 15.3 27 43 534 (i-4) (+4,5) (8) (±8) (4,7) 4 4.2 6.3 (±1.1) (±2.4) (+) 7 8,0 13.8 15 31 (+3.7) (+í.) (+11) (±8) Petticce o 7.3 37 88 114 117 Wick 1.6) (+10) (+io) (+21) (6) Bay 1 6 45 76 106 123 130 (ii) () (+6) (+ii) (+io) () 9 38.5 77 64 (+io) (-i.,i) () 11 1.4 13.6 20 28 29 36 (+0.8) (±4.2) (±7) (+6) ) (6) FI}Th/S8Laminaria hyporborea

A D

D n V ot o V o

D D

u A o n ri t

i 2 6 7 8 9 IO II 1213 4 Number nlrings Fig. 12Measurements of individual whole plants oollec'ted from thrououteach a ling area with different numbers of' rings in the stipe (approximating to age). A, Mean fresh weight. B. Mean length. , Kvitsy Kariiy £ 9 4>, Nord Frya 3. Helgeland 0 E. Finrk Pbr site positions see Table IX,(Data replotted from Grenagex', 1953, 1954, 1955, 1956 and 1964.) 4:8 FRr4/387 Laminaria hyperborea At Ytteraiden (Baardeeih, 1954) the lengths retrieving whole plants.The grab used 'by were 1.6 a (3.4).It seems likely that the Walker in Scotland sampled half a square yard maximum length of this plant is reached only and that usad by Grenager in Norway half a between the latitudes of about 55°N and 65°N. square metre (loo x 50 cm or 71 x 71 cm). It is Rain (1967) concluded that southwest Norway is olear that this method must miss some of the at about the optimum latitude for -the species. biomasa and some of the individuals.It may also As has been shown, there is little effect of sample relatively blind.It may land on unsui- latitude on the size of the plant at any given table substrata.In expressing density or and the greater maximum size at the optimum standing crop from this method there is a choice latitude is mainly due to greater longevity of averaging the results from all the samples or (iain, 1963, 1967), only those which oontained weed.The first alternative would give an artificially low In most cases, although the stipe varies in result on a mixed bottom if density on rook is appearance under different conditions it does not required, while the second alternative may give vary in basic shape.There is a straight line a high result in deep water where density is low relationship between the logarithm of the stipe even on suitable rock. weight and the logarithm of the stipe length whlch is the same for most populations from very dif- Densities, in terms of numbers of individual ferent habitats which include deep water and thus plants per square metre, obtained by various low irradiance (Rain, 1971).Figure 13A shows authors, either through -the use of -the spring this relationship and incorporates the data from grab or by diving, are shown In Table VI.It la five populations.There seem to be, however, at once clear thai the results obtained by diving occasional e:ceptions to this relationship.In are considerably higher than those obtained by some sites, John (1968) found stipes which were -the spring grab.This would be expected from abnormally long for their weight.Figure 13B shows the difference in technique.When all -the young plots of ag group means of' ash-free dry weight plants can be collected by a diver the apparent against lenE'tb from a number of sites (John, 1968).density must be more variable and particularly Sorne from polluted water (squares) can be seen to high wIzen conditions have allowed a plentiful be off the line -to the right, particularly in growth of aporelinga.The grab data, based on large plants.Here stipes 2 m long have -the saine many samples and including mainly mature plantr, we:Lght as stipes iin long from moat other popula- are much more consistent. tions.However, two further populations showed similarly long lengths at these weights, those LUning (1969a) made use of a concept from off the island of Coil in the inner Hebrides initiated by hiher plant ecologists, the leaf (triangles),Here the water was clear and unpol- area index (LAI), or area of leaf per unit area luted.What caijses -this effect is thus obscure. of substratum.He found an index of 41 on a There is no evidence that it is due -to "e-tiolation" bottom a-t 2 n below mean low water springs, as suggested by Bellarny and Whlttiok (1968) 2.3t 3 n and 1.6 at 4 mThe last were thus because it is not shown in deep water (Rain, 1971a), aproaehing an open community where there is 4.13Sporophyie-gainetophyte and sex little self shading.Drach (1949) gave figureo composition equivalent to 6 -to 8 for populations in the Phglish Channel. Nothing is biown of the density of gameto- phrtes in the field bu-t i-t is likely that they 4.22Variation in density with normally far outnumber the eporophy-tes which in ecological and accidental fac- turn are astronomically greater in biomasa. tors Schreiber (1930) isolated mingle unilocular The moat obvious factor affecting density sporangia and found that equal numbers of males is depth,It is apparent from all the obser- and females developed from each,Unless their vations made a-t different depths in Table IV aux-viral is differentially affected, therefore, -that there is a progressive decrease in density the sex ratio of the gametophy'bes is 1:1. in deeper water.This is presumably due to decrease in irradiance, and is more rapid in 4.2Density turbid water such as that around Helgoland. Ernst (1966) found that the total density of a 4.21Average densities of defined mixture of L. digitata, L. hyperborea and L. areas ochroleuoa in north Brittany decreascii logarith- mically with depth,It ham been suggested Nos-t of' the surveys of populations of L. (Llining, 1969a;Rain, 1971a) that the decrease perborea have been made by the spring grab in density, and thus in self-shading, with depth technique.In this a grab is lowered from a partly couriteracts the decrease in irradiamos boat end uhen protruding feet touch the bottom and this results in the amount of li.t reaching -the jaws aro closed by powerful springs, the the plants below the canopy being relatively teeth of the jaws either cutting atipes, usually constant with depth, within limits.This could near the rock, or pulling holdfasta off, thus explain the relatively small change in apparent FIr/87Larní.nEria hyperborea

01 01

Leng h of st in - fl Length of stipe -- m .g0 13A, Stipe fresh weights plotted against stipe length for populationa ,tE4, HE17, BE33, BS1ß (see Table IX for positions).(Data mainly fromKam,1971a0) B, 5-tipe ashfree dry weit plotted against stipe length for the following population , BbAbb's Head. 55°54-5'N, 2°ß-9'W Be..ell 5°33'N1°37'W; Saddell Bay 55°32'N, 5°30'W;Rubh'n Amair 55 46'N, 5°5'W;Cardigan Island 52°8'N, 4°42'W;and Sennen Cove 50°5'N, 5°41'W. ,tarsden 54°58'N, 121'W and Redoar 55°38'N1°2'L. ACoil 56°34'N, 7°39'W, (Data frein John, 1968e,) 10

TABLEVi

Density, as number of plante of L0 hyperborea per square metre at various depths sampledbyWalker (i), 1950, 1952, 1957; Walker and Richardson (2), 1955, 1957; Gren:er (3), 1953, 1954, 1955, 1956, 1958 1964; Bellamy et ai.. (4), 1968; John (5), 1968; Whittick (6), 1969; LUningb),1969a; and iin (8) unpublished. For latitudesandlongitudes see Table IX.

Plants/rn2 in samples with weed Meen-sil depths Locality Year With All weed samples

Obtained by grab:

Oirven (2) 1952 7.8 1953 8.6 7.0 5.5 2.4 1954 9.4 9.9 6.5 1955 9.1 8.3 5.2 Aliso Craig (i) 7.4 6.6 Dunbar (i) 1952 5.6 5.8 4.3 3.4 5.2 (2) 1954 9.1 7.3 4.6 3.2 (2) 1955 8.8 7.7 6.1 4.6 NEMay Island (i) 16.3 10.5 9.0 8.0 serburgh (2) 1952 4.8 6.2 6.0 4.4 3.2 5.6

1953 6,2 7,5 6.0 4.5 3.0 1954 7.0 6.8 6.2 5.4 4.5 1955 7.6. 9.0 9.1 6.8 5,5 Kv-itsy (3) 9.3 8.6 Kaniy (3) 9.6 8.8

Pus na (3) 8.3 7.9 Sub (3) 9.3 Froyene () 8.4 7.8 S.Helgeland (3) 7.7 7.5 Hely (3) 5.7 3.4 E. Finrnnrk (3) 6.1 2.7

Obtained by diving:

SeimenCove(4) 27 3B 42 12 Helgoland (7) 27 12

Piombo rough (6) 16 10 5 leleofMan (8) 27 39

PstticoeWick Bay(5) 34 23

(6) 20 23 17 6 Connel Sound (8) 1970 23 17

Muidoanich Is. (8) 1970 46 14 Off Loch Boisdale(8) 1970 0,2 1/S87 Laminari a hyjperhorna rowth rate with depth under forest conditions is greater where there is water movement (see (see 4.12), 1.31), and the size and extent of the hold-fast increases with the size of the plant, there must Another factor affecting density mey be come a stage in the development of a plant latitude.There is no general trend apparent ( except uider very sheltered conditions) when from Table IV but it is possible that densities -the attachment is inadequate to hold its bulk are lower near the northern limit of the species, against the movement of the water,This explains particularly if all the samples and not just why plants at 1 a in the Isle of Man have a lower those with weed are considered,This is suppor- longevity than those a-t 2 m or more, because wave ted by Kath's (196'() observation that apparently action is most severe near The water surface. suitable substrata are weakly colonized by L. Anything that decreases the efficiency of the hyperhorea in north Norway.One woult expect attachment mechan em mu st increase mortality. this of any plant near its geographical limit Kitching (1937) concluded that just above low but it would be unwise at present te guess the water springs attachment to barnacles instead critical factor involved. of rock surface subsequently led to breaking away, but this is unlikely to be important in There is evidence -that in some areas deeper water because barnacle cover is less grazing can affect the density of this species. dense.In much of Britain a serious cause of Jones and }(ain (1967) regularly cleared Echthus weakening of the hold-fast is grazing by Patina eculentu,s off an area of rock at 10 m below pellucid-a which eats cavities in -the base of the LiT normally bare of lamLtharians and eri algal stipe and detaches it from many of the haptex'a. population developed which after two years The chances of infestation seem to vary from included a vean density of L. hyperborea of 4.4 place -to place,In the Isle of Man they are plents/squa.'o metre, high, with all age grOUpEs above on&.-year oldie affected, and at i a below LiT the mean infea- 4.34irtality, morbidit tstion of five to aeven-'ear old-s was 54 percen-t (Kath arid Svendsen, 1969).In ßerwick, south- The maximum ago of plants in the Isle of east Scotland, however, in a- large sample from Nan so far found by ICain is ten years.In the o io 5 m no two or thro-year old-c were infested Norwegian populations studied by Kam(1971a) arid -the nican infestation in the same age range the maximum age was li years but larger samples was only 14 percent (1hi-ttick, 1969),in both could have revealed older plants.Grenagar cases infestation increased with açe up. to scrc certainly found plants with more rings though9 years arid -then dropped, suggesting an efteet on as aireedy mentioned, these may not be very mortality,Many examples of brohen hcldfaita reliable as indications of a-ge.However, as nithout stipes with Patina cavities bave been - some of his plants were very large and his seen in the Isle of Man,lloweveralthough sanples numerous, it is possible that all the Kit-'-ting (1937) found that 90 percent of the L. rings represented. years.The highest was 18 borea plan-ta washed up on the beach near in E, Finmark (Grenager, 1956),He also Plymouth (southwest 1igland) after storms had recorded 16 in Nord-]rya, 15 iii Jjssund been detached- et the holdfnst, only a few of (Grenager, 1964), 15 in S. Helgeland (Grenager, -them were infested with Patina.This, thereforo, 1955), 14 in Tustna (Grenagex', 1954), 13 in seems an area of lower risk then the Isle of Nan Kartry and 12 in Kvitsjy (Grenager, 1953). John (1968) also found relatively low infesta- -tion levels in sites in northeast and southwest This perennial species shows no obvious Thgland and west Scotland, with a mean of eight mechanism for the decay of any part of -the indi-. percent in five to seven-year old-s.ifowever, vidual or for obvious senility of the whole, his data included all depths and sites.There The old frond is shed after about a year but je evidence that infestatIon decreases uith -this is probably a mechanical process rather depth (Vaiü, 1971),it is known that there is than a progressive degeneration as in some a change in behaviour of the limpet with lati- species of the genus,Nor does -the whole plazrL tude, In France (Sauvageeu, 1918), souhwest cease its capability for growth after a limited flegland (Graham and Fret-ter, 1947), the Isle of period as do soma genera in the order, The Nan (Kein, 1963) and sonthoao-t Scotland capacity for growth may be reduced after some (Whittick, 1969) i-t penei;rates -the hold-fast bu-t years but as long as the plant remains intact, it does no-t in parts of west Scotland (Kein, attached and -the conditions remain suitable, new unpublished) or Norway (Hain and Svendaen, 1969). -tissue seems to be produced each growing season, Phi s may partly explain the apparent great e r longevity of L, ea in Norway than in the Amongst outside factors the most obvious Isle of Man. cause of death is physical removal of a plant from the substratum,The force causing this Before the holdfast becomes inadequate to removal is likely to be wave action or other hold a plant in the prevailing conditions some water movement, ihile that resisting i-t is the other accident may have befallen it,Possibly power of adbesion of. the hold-fast in relation the most likely is grazing by an echinoid,Near to the size of the plant.Although this adhesion their area cleared of Echinun osculentus, Jones :12 FIR?4 S87 Laminaria hyperborea and Kam(1967) found that, although in July miss small plants than large, moro difference sporelingu of L. 1ypex'borea had developed on would be expected between the two methods in boulders at 10 s up to a density of 20 pmants/ terms of numbers than of weights..However, when square metre, by November almost all had disap- Gronager (1956) and Baardseth (1954) cleared peared except where clearance continued.Thus quadrata by hand they obtaind 1.5 to 3 timos Echi.nuwas responsible for the destruction of as much fresh weight in k/m' as obtained by the ninny spOreling3 of less than 100 mii.It is likely grab technique. also that Echinus can damage mature plants irreparably.The animals are frequently to be In most cases9 as expected, there in a seen grazing on the plants or their epiphytes, decrease of standing crop with depth (Table Vii) and when this is ex-tensjvo at the transition zone and ilma with irradiance.Walker (1954) stated recovery may be impossible.Patina may also that this decrease was sore the result of a damage this vulnerable region (Kath and Svendsen, reduction in density (individuals/unit area) 1969).However, stipes are frequently found than of weight of individual plants.This was broken farther down and the reason for this is confirmed, by LUning (1969a).This does not mean9 not apparent though it can be caused by the plant of course, that there is no decrease in plant becoming entangled and the stipe bent sharply for size with depth, because this has been shown to a period.On fertile fronds Patina frequently be the case in some instances (Kam, 1963, 1967; grazes on the sorun, destroying the eporangia, LUning, 1970a;John, 1968;Whittiok, 1969) but Another graser which may be important in Norway this is unimportant frein the point of view of is Lacuna vincta (Vail, unpublished).Vahl found standing crop, compared with density decrease, considerably higher densities of this gastropoci than of Patina at Lyrrodane in Norway and the Considerable differences were found by holes it muios are deeper. Walker when he repeated some surveys in areas in Scotland,In one case, at Dunbar, this was It seeiis unlikely that fronds are regularly probably due to accumulation of sand (Walker grazed by f:.sh,One sparse population of plants and Richardson, 1955).Taking the surveys shown wan found iii the Isle of Nan with their fronds in Table VII, which included only L. byperborea partly eaten (Kam, unpublished).This was in and in addition further surveys and their a regioñ of fairly fast tidal stream and. sub- repeats includinalso L.jtata and. sequent observation showed that in many cases the charma, Walker (1956a) found that there was a fish Labnis berylta was stationed dounatream of generaltrend in standing crop from a peak in each plant apparently using it as cove'0The 1947 to a trough in 1953,This he later cor- fish snapped at food passing in the current and, related with sunspot activity (1956b).There because the frond was waving about in foni of has been no confirmation of this somewhat it, occasionally took a bite out of it.This unlikely possibility. sort of dunage is unlikely to be important in mortality, The total standing crop of stretches nf coast iherç L, hyperborea was almost exclusively Diseased plants have sometimes been observed, dominant is shown in Table VIII,Chapman's Fronds of the population B54 in Norway (Kam, (1948) figures were based on a somewhat hurried 1971a) were blackened, at their tips and were war-time survey using a grapnel and echo-sounder decaying but the causative agent wan not identi- from a boat end aerial photography.Walker and fi. ed. Richardson's and Granager's figures were the result of surveys with the spring grab previoualy 4,4 Total quantities (standing crop) of mentioned.Boney (1965) compared Walker' s and defined areas Cpms data for total Laminariaoeae and oon- Surveys of the standing crop of L. hyperborea eluded that more detailed mapping results in have been made almost exclusively using the spring higher estimates.In his surveys Walker usually grab (see 4,21).Unfortunately Walker always applied the following formula to his results and used the term "density1' for the fresh weight per obtained values for k andtaccording -to the unit area instead of its more usual use for the site: number of individuals per unit area,Again there iS a choice of expressing the results using all IT the samples or only those which contained weed. c kf In Table VII the data haveheon selected or recal- where d fresh weight of weed per unit culated from samples with weed, this being more area in all samples, realistic in shallow water but perhaps less in deeper water.The only samples in the table C % samples containing weed, obtaiued.by diving are those of LUning frein f depth. Helgoland.They are comparable to the grab sam- plea instead of being much higher as were his (Walker and Richardson, 1955.) density figures.An a grab is more likely to FI11/S87 Laxninaria hperhorea

TABLE VII Standing crop, as kilograznnie of fresh weight of L. hyperborea per square metre at various depths sanpied by Walker (i), 1950, 1952, 1957;Walker and Richardson (2), 1957 Orena«er (3), 1953, 1954, 1955, 1956, 1958, 1964;Baardaoth (4), 1954;and Lilning (5h 1969a. See Table IX for thtitudes and longitudes.

kg/rn2 in samples with weed Locality Year 0-2m2-6m6-10m1-14m14-18m18-22m 22-26m26-30m Obtained by grab: Giivan (2) 1952 8.0 7.0 1953 6.2 5.3 1954 5.3 5.3 1955 5.5 5.0 Misa Craig (i) 1955 5.7 3,5 Dunbar (i) 1949 6.7 5.33.4 0.9 (2) 1952 3.6 3.4 2.1 1.1 (2) 1954 5.0 3.62.0 1,0 (2) 1955 5.04.2 1.8 1.0 ME May Island (i) 1956 5.5 2.9 1.7 0.9 0,8 P'raserburgh (2) 1952 3.3 4.5 4.0 2.2 1.0 0.7 1953 3.1 3.6 2,9 1,6 0.9 0.8 1954 4.44.3 3.3 2.7 1.8 2.0 1955 3.4 3.8 3.2 2.2 1.3 1,2 Holm Bound, Orkney (i) 1948 5.9 5.64.4 3.4 Sanday Sound, Orkney (i) 1947 7.3 7.0 6,1 3.8 E. Sanday, Orkney (i) 1948 7.1 7.7 7.3 6.4 N. Saudny, Orkney (i) 1948 7.1 7.7 Kvitapy (3) 1952 8.8 5.2 3,6 3.4 1.0 0.2 Karry (3) 1952 7.6 9.8 7.8 4.2 4.2 0 0 Tuatna (3) 1952-3 .8.6 9.09.0 6,0 5,4 3,4 1.9 1.2 Buia (3) 195412.6 10.010,0 5.0 Fropene (3) 195412.0 9.2 6.8 4.8 1.2 0.6 S. Helgeland (3) 1952-39.0 9.08.2 4.4 4.2 1i (4) 1952 6,9 4.3 Karlsyvaer (4) 1952 5.4 5.9, 3.5 3.9 1.1 Nordlandaflaket (4) 1952 5.2 2.0 3.6 2.2 2.0 N. MoEkeneey (4) 1952 6.8 7.8 7.4 3.4 2.7 1.6 E. Noakeneey (4) 1952 6,1 6,85.2 3.5 2.4 1.7 0,6 Viiken (4) 1952 3,5 3,0 3.1 2.2 1,5 Hely (3) 1953 7.4 6,0 4.8 3.6 1.9 1.7 0.2 E. Finmark (3) 19535.167.38 6.1 3.88 2.4 2.4 0.2 Obtained by diving helgoland (5) 8.7 4.7 4:14 F]FS8Laminaria erborea

TABLE VIII Estimated standing crop of weed on surveyed sections of coast dominated by L. hyperborea at all depths.Da-ta from Chapman (i) 1948;Walker (2), 1950, 1952;Walkor and Richardson (3), 1955;Grenager (4), 1953, 1954, 1955, 1958, 1964;and Baardseth (5), 1954. See Table IX for latj-tudes and longitudes.

Standing crop Locality Ar kin' Metrjo tons

Berwick/Newbiggin (1) 119 600 Dunbar (2) 13.8 51 400 St. Anth'ews (i) 43 800 lintore (i) 53 200 Halmsrb.le (-t) 60 600 Bound of rra/W. Loch Tan-bot (i) 682 000 Fraeerburgh (3) 2.93 11 700 Holm Sound, Orkney (2) 5,3 19 800 Banday Sound, Orkney (2) 26.3 107 200 E. Sanday, Orkney (2) 19.4 129 000 Kvitsy (4) 9.7 65 000 Karsy (4) 10.6 76 000 Tuetna (4) 7.0 60 000 Bula (4) 25 250 000 Proene (4) 20 140 000 S. Helgeland. (4) 49,3 312 000 W. Moskenesy (5) 3. 94 14 000 E, Moekeney (5) 1.96 7 000 Vikien (5) 22.25 46 000 Hely 100.6 281 000

4.5Accompanying species of the hglish Channel L. saccharina may dominate as it does also on unstable boulders which Over most of i-te geographical range L. may be turned over without damage to its hyperborea is the longest lived dominant species., flexible s-tipe.From the Fhglish Channel south- This explains its success in comp-ition with ward L. ochroleuca is a serious competitor in fas-ter growing cxaniples of Lainina::iales, Once slightly sheltered sites,This species, similar l-t has become es-tablished the high canopy foriaed in habit -to L. perborea, is co-dominant with by the fronds effectively excludes almost al). it in many placeB where their geographical serious competition.The species with which it ranges overlap (John, 1969) and replaces i-t in competes for establishment are all members of shallow water in Spain (Scanne Gamba, private the Laaninariales,In sites with very severe communication).Finally, L. hyperboLea com- wave action Alaria esculenta is successful in petes with opportunists in situations where the shallow water. substratum is temporarily denuded.The most successful of these south of Norway is Saccor- Juet above low water sprin where wave hiza polyschides.Other brown algae ouch as action and exposure to air are not -too severe, Desmarestia viridis, D. aculoata and even Alaria i-t is L. digitata which succeeds in place of esculenta and Laminaria saccliarina can act us L. hyperborea,Under sheltered conditions north opportunists and dominate for a period in sitas /s6Laminaria ,erborea

TABLE IX

Latitudeaand.longitudea of sitee in prOVIOUB tableo

Sito Latitude Longitude Sito Latitude Longitude

Ailoa Craig 55°16'N 5°7'W Li O 54°3.5'N 4°46-7'W Balintore 57°15'N 3°55'W LooL Boiudale 57°9'N 7°15'W

BCuO.5 60°19'N 5°16'E !faybland 56012e N 20321W BE4 60°9'N 5°7'E N0a4 54°7'N 4°44'W BE33 60°9'N 5°7'E ME4 54°5'N 4°46'W 5403 13erwick/ 55°47'N 2°W NEI4 4°47'W Newbigin 55°12'N 1°30'W 54°3'N 4°46iW 6O°10'N ZH14 4°59'E NS4 54°6'N 4°45'W 60°10'N 5°E Muicloanich bland. 56°55'N 7°26'W Bs18 6o°16'N 5°13'E Nor landaflakot 67°45'N i 2°45'E Corinel Sound 56°28'N 5°24'W N. Sanday 59°18'N 2°32'W Duxibaz 55°57'N 2°24'W Petticoe Wick Bay 55°55'N 2°9'W Dunmanu Bay 9°45'W 51°35'N Reine 68°O'N 13°O'E E. Finmark 70°21'N 31°7'E R,et 67°30'N 12° 10' E E. Noekenee$y 13°10'E 67°58'N t. Andrew a 56°20'N 2 18' W E. $anday 59°16'N 2°30'W Sanday Sound. 59°12'N 2°40'W Eepevaoi' 59°30'N 5°6E SE6 57°44'N 5047'lf F2 54O4...5tN 4°46.7'W 5E10 57°44' N 5°48 'W 4°46-Î'W 5707 54°3-.5'N $E16 'N 5°48'W Flanborough 0°5'W 54°8'N Sannen Cove 50°5'N 5°25'W Fraeerburgh 57°42'N 2°4'W 8,hieigoland 65°25'N 1 E yene 64°2'N 6°54'E Sound. of Baz'ra/ 7°25'W Girvari 55°10'N 4°57'W W. Loch Tarbot 57 55'N 7°W helgoland. 54°10'W $85 55°42'N 4°57'W Uely 70°12'Zl 18°0'E Sum 63°0'K 8°25'E Helinedale 5807'N 3°40'W 69°3&-55 'hi 18° 1-43 E hole Sound. 58°52'N 2052'W Trendheiaafjord 63° 30' N 10030, E lele of Man 54°5'N 4°46'N 63°1 t 'N 7°55'E Karleyvaer 67°33'N 14°50'E Van1S 70°20'N 31°O'E Eaxy 59°16'N Vitken 68°58'N i 3° 15' E Kvitepy 59°3'N 5025'E W. Moakeneay 67°57' N i 2°55'E L5 54°4-5'N 4°46-7'W 4: i6 /8ï Laminaria hyperborea where the climax vegetation is dominated by water and the distal part of the stipe while L. hyperborea. Phycodrys rubens was more important in deep water.Ptibo-ta plumosa and Membranoptera alata The algal r;pecies forming the undergrowth were mixed with or slightly distal to Phyc[s. to an L. .h.yperborea forest vary somewhat with On the other hand LUning (unpublished) reports the iatjtudo,On the French coast of the Ehglinh that Polysiphonia urceolata and Membranoptera Channel, the Channel and Scilly Isles, the fol- alata are the only abundant stipe epiphy-tes in lowing species are common in the undergrowth Helgoland.Whittick (1969) found that the (Drach, 1949;Ernst, 1955;Kam, 1961;Norton, epiphy-te biomass was greatest at 1 is below L.kf, 1968);Calloph,yllis Ïacjnjata, Delesseria showed a seasonal peak in September and snnguinea, Crrptopleura raxnosa, Phycodrys rubens, increased with the age of the host up to five Pio canium oartilafjneum, Pteroaiphonia parasiti ca, years.Whittick (1969) also observed sites in Diotyota &Lchotoma, Phyllophora crispa, Kally- Yorkshire, southwest Ireland and extreme south- mania reniformis, Dilsea carnosa, Heterosjphonia west Ehgland..In all Ptilota was absent and plumosa, Halopterta filicina and Dictyopteris Rhodyinenia extended deeper.In southwest membranacea.Though none of these species is 1gland there were no epiphy-teo below 22 n absent from the Isle of Man only the first eight although L. hyperborea extended to 36 m, are common in the undergrowth there, but ad&i- Kitching Ç1941) found Dolesseria sanuinea, tional common species are Öutieria multifida Odonthalia dentata and Cryptopleura ramona sporophyte, Odonthalia dentata and Desnurestja abundant on stipes in west Scotland and Marshall aculcata (Kam, la), All these species are (1960) also found that Plocamiurn cartilagineuin, also common in west Scotland (cAllisier et al,, Pterosiphonia.rasitica, Calbophyllis laciniata, 1967),Missing from this list in Helgoland Ptilothauinion pluma and Plumaria elegans fairly are Calloph].ljs, Cxyto leura, Pterosiphonia, common in southeast Scotland,Eoept for Cutleria'ant OdonthaliaLU.nimg, 1970b),How-' Odonthalla, Cryptopleura, Ptilothannion and ever, a numer of other species are relatively Plumaria all these species (in addition to more common yllophora membranifolia, othFwsre found on stipes in Nt-Fya Corallina officinalis, Ulva sp.In southwest (64°N) in Norway (Grenager, 1964).On the other Îlorway the only absentee of the species common hand, .Jaasund (1965) lisis as epiphy-tes only in the Ile of Man io Cryptopleura,However, Rhod,ymenia palmata, Polysiphonia urecola-ta, neither Plocamiurn nor Pterosi honia are common Odonthalia denta-ta, 1h-thora cris-tataon haptera there (Breivik, 1958; In north Ectocarpus fasciculatuÇn fronds) and Norway most of those species are absent but Phloeoapora cur-taFbshie) Jaasund (on fronds). Pbycoth's, Ptilota plumosa, Deamarestia aculeal;a E. Í'ascioulatus is also common ori old fronds in and D, viridis are abundantJaasund, 1965). the Isle of Man (1hssell, 1966), along with Delesseria and Odonthalia are also present. Giffordia hincksiae (Thisaell, private communication) and Laminariocolax tomentosoidea Drach (1949) estimated that in an L. (Bui311, 1964). hyperborca forest of a density of 20 plants per square metre, the area suitable for settlement, Sloane et al. (1957) investigated the provided by the rugose stipes of this species, animals inhabiting the fronds of L. hyperborea could be 150 percent of the rock surface.This near Lough Ine in southwest Ireland,The additional habitat is particularly favourable commonest was Patina luida, followed by to plants because they receive more light than Nembranipora membranacea and Obolia Lenículata. on the rock surface.Tokida (1960) lists Also recorded were Hippothoa hyalina, Sertularia records of epiphLy-tes on L,Tperborea Stipes operculata, Scruparia clislata, Hiat ella arcti ca and fronds totalling 71 species.Many of thea and anomiida, are chance inhabitors and the main occupants of thetipoa are fairly constant in any one boar- The holdfast of L. erborea forms a home lity.Marshall (1960) and. WKittick (1969) to a great many animals, far too numerous io studied the epiphy-tes of L. hyperborca stipes lint.For example, Moore (1971 a) found 387 in Berwick, southeast Scotland and both found species in holcìfasta collected only from the that four opecies were of particular importance. northeast coast of Britain from Berwick to Mhodymenia palmata was confined to shallow Yorkshire.Of 61 species of nematodes, five were regularly abundant (Moore, 1971b). FIF4J337 Laminaria hyporborea

5 kUR1flSTING O and 5 m and had a capacity of 380 kg (fresh weight).A much larger dredge ("trawl"), collec- 5.1Equipment and methods of harvetin ting 1 000 kg, with a horizontal knife, is now in regular use in Norway. The simplest method óf harvesting, which ham been used for centuries, is the collection A number of experimental approaches to the of cast weed.This je the only method ucd in problem of drying L. hyperborea have been made. Britain now. Sjckles have sometimes been used About half the water was found to be removable to cut weed at low water, without heating by Reid and Jackson (1956).The mechanical methods which they employed were Grapnels can be used from boats to haul in centrifuging after hammer-milling, batch pres- plants of L. hyerborea.The normal boat grap- sing, squeeze rolling and screw expelling. nei with prongs curving inward from the end, of Detailed inveatigationa have been made of ther-- the shaft js less effective than a specially mal through-circulation drying of the stipe deaied instruisent with prongs curving outward (Gardner and Mitchell, 1953a) and the frond. from an acute angle (Chapman, 1944), Chapman (Gardner and flitchell, 1953b).The material had found that a diamond-shaped instrument was to be cut fairly fine for this method to be better still.In Scotland, at what was later economical.The design of through-circulation called -the Instituto of Seaweed Research, dryers was summarized by Gardner and Mitchell another type was developed but found inadequate (1956).chat was claimed to be a more economical for commercial harvesting (Jackson, 1942). method of drying this species was described by Booth (1956)..J1illed fresh material is poured In further work at the Institut ethe between two parallel steam-heated drums rotating continuousrapnel (Hay, 1952), later called 'the in opposite directions in such a way that it is belt haryeser (Jackson and Wolff, 1955; squeezed in-to a thin layer as it moved downward Jackson, 1957), was developed,This consisted of between the two drums, adheres to each, is dried a conveyor belt of flexible wire mesh and chaina quickly on thesurfaces end is scraped off at the edgeB with rows of hooks across at inter- before a full revolution of the &rwa takes place. vals.The weed was compacted by the mesh, torn off by the hooks end carried up to the boat on In practice L,yperborea is treated in the belt.This system was tried out extensively Britain as follows.Stipes cast up on the shore and seemed. to be fairly satisfactory though its are colleoted and air-dried. locally in stacks, depth range is clearly limited.It is not in Fronds are no-t used..The half-dried stipbe are commercial use. sent to South Uist (Outer Hebrides, Scotland) where they are further dried in batches on iron Another method. on which the Institute -tip-wagons in a heating chamber through which worked consisted of using a cutting heaLi bot air is circulated,The material is then (Jackson and Molver, 1952) in the weed bed and mil1 ed -to a mea]. and sent to an alginate fac-tory mucking it up to the boat in a flow of water in west Scotland. (Jackson and Macduff, 1952).They had difficulty in controlling the entry of the weed into the In Norway, on the other hand, this species pipe and although this problem was solved in a ja actively harvested in situ.Nest of the test tank (Jackson, 1957), the developed suction material is collected from Í'ishing vessels or method was never finally tried out in the sea from specially designed boats, using a dredge and is not used commercially. ("trawl") as already described.The weed is cut off above the holdfast by the knife at the The only apparatus in commercial use is the leading lower edge arid. the plants pass into the "trawl"This term is incorrect because the rigid box and accumulate at the back.Nhen full apparatus has a rigid lower boundary to the mouth 'the dredge is hauled up and emptied into the arid should thus be called a dredge (Davis, i958). boat.Either the material is sent to a factory One of the earliest of these was used by the within hours or it is chopped, mixed with for-' Institute of Seaweed Research to sec'ure nearly malin and acid and. packed into a silo, of which whole plants (Jackson, 1949).I-t consisted of a number are distributed along the Norwegian an open-ended rigid box with rows of triangular coast.The material is later transported to knives at the mouth,It was used little because the alginate factory in southwest Norway. the knives became clogged.Another dredge, which was used commercially for some years in 5.2 Harvesting seasons Norway, is described byin5rholm (1964) end Svendeen (in preparation).This also consisted In Britain, because only cast-up stipes of an open-ended box 1.5 a long, its floor being are used, harvesting is mainly in the winter a plate of steel and the sides and roof of wire when there are nost gales and thus strong wave mesh with a row of triangular knives at the action removing plante from the rock. lower edge of the ¡south.It was set on runners to raise the knives off the bottom and had a In Norway dredging takes place throughout float on the wire ease,It worked best between the year but, again because of strong winds in /S37 Leatnaria hyperborea winter (lu this caso unfavourahie), the best holdfaats and ziïiall planta intact.The standing seasons are spring, Sumner arid early autwrn. crop (fresh weight) after 2.3, 4, 6 and 7 years was reapectively 6 to 1O, 12 to 23,15 to 31, 5,3ilarvestinareas and, de the 25 to 38 and 22 to 33 k/m',An he observed values of 20 to 50 kg/rnin virgin forest the British supplies como from west Ireland standing crop bad returned to its original level (flood, 1953),the Outer Hebrides, 'the Orkney in about five years.However, a mazinein stipe and the Shetland Islands.Presumably planto length of 2,44¡uwan observed in the forest and from aU depths within its range of growth are a length of 1,4 'to 2.3 n was only attained after included in the casi, six yearn on the harvested. areas,This rate of growth was very fast. under these optimal condi- In Nor;oy harvesting talcou0placc on the tions where competition from opportunists seems west coast betueen Jaren (ca59N) end Trond,el to be less -then in the Isle of Pian, (ca 65°N).Soiie hand collection takes place down to i m below low water,A small dredge io 5,5Total annual elda used from 1 to 5 n and the larger one from5 'to 15 n, Because of the secrecy adopted by the algina-te producers precise data on the total 5.4Regr'ow-th on harvested areas uan'tities harvested are not available.It io probable that in Britain and. Norway together a The first direct observation ori rogrow'th total of just under 4 000 tone of alginate ax was e by Kitohing (1941),He out forest with produced. from L. J,jajo rea each year.An the shears and. found 'that one year later the renal'- weight of aigine.te produced is about 6 percent ning holdfasts had disappeared. and there was a of the original fresh weight of the algae, this dense growth of new plants of L, hypex'borea with corresponds to a total annua], ex-op of about a total height of about 1¡u. 60 000 tons of fresh L. oexorea,Clearly this varies from year 'to year, partloularly in Svendsen (unpublIshed) has observed an area Britain where i-t depends on the amount cant up0 in Norway off Vevang (63°fl, 7°'16'12"E) which had Based on Norwegian prices in 1971, tIja total been harvested at a depth of 2 to 5 n with the value of this orop at first hand crreapondn to saaweea dredge ("-trawl", see,i), which leaves about £200 000, 6 PROPXTION AN]) 14ANACthIWf 6.5Artificial culture 6.2Control of physical factors In laboratory culture work the emission of zoosporea from fertile fronds of L. b,'y-erborea is The only obv-ious way to creato a habitat fairly easily induced.A fertile frond, profer'- suitable for L. hyperborea is to provide a sui- ably with some sien of dehisoence at the tips table solid substratum where none existed before. (greenish patches) is removed from the seas An accidental example of this is Port krin bresic- drained and. put strai..t into a polythene b water in the 181e of l'ian.This was built in the which is left overnight in a cool placo.Suit- nineteenth century on sand ai about 10 m below able fertile parts of the frond are then out up LAT and was soon pushed over in a storm.It is and placed in sea water.If emission &)es not now nn untidy strip of concrete blocks and boul- occur within 30 minutes it may be stimulated by dera 230nilong and 70niwide, rising from 10ni a rise in temperature but not above 200G, io about midtide.It forms an winirable substratum for L. hyperborea over much of its area, Iflning has used two methods of transplanting where before there was only sand.. this species for experimental purposes.In one (LUning, 1970a) he left polypropylene ropes 6.4Control of biological features attached to iron frames near L.yperborea plants in January,After some months sporophy-tes had 6.41Control of predation and developed on the ropes which were cut into pieces competition and attached to P.V.C. plates mounted on an iron frame,The new haptera of the al:e soon One of the moat serious predators of L. attached them to the plaies.In another method hyperborea is Echinus.This can be removed or (L4lning, 1969b) he carefully removed plants destroyed by hand by divers but this is a already growing on the rock, placed them on tedious and costly process.The method used for P.V.C. plates and held them there with a nylon sea urchin destruction in the programme to improve network and. rubber bands.Outgrowths of the the Nacrocystis habitat is the iissemination of existing haptera soon attached the plants and the quicklime, CaO (Leigh-ton et al., 1966).It. is network and bands could be removed before the claimed that eohinoids are the most sensitive outgrow-th of new haptera above them. organisms to this substance,A concentration of 0.5 kg/rn2 is allowed to trickle from the stern Svendsen and Kein (1971) also transplanted of a boat moving along a grid and the chemical individuals from one site to another.The hold--- is distributed by the propeller turbulence,The fast was removed from rock and lashed to a stone particles should be large so that they sink with thin nylon rope.The stone was placed in ç. quickly and make contact with the urchin before bed. of cement mixed with rapid. hardener which had the reaction with water is completed. been taken underwater in a polythene b, There is at present no known method of John (1968) transplanted first year plants controlling predation by Patina pellucida. on to underwater frames by enolosing their hold--- fasts in longitudinal slits in poly-thene tubing and binding the tubing with thin poly-thene rope as close to the holdfaai as possible. FIRI.i/587 Laminarir hyperborea :1

7 CE»tICAL CC)MPOUITIUN water, fresh weighi can clearly be variable according to the tochniciue used to determino it. In common with other species of the genus, To avoid drying out of the plant, it should be L. hyjorborea has a high content of carbohydrates removed from sea water immediately prior to and mineral matter,There are appreciable dif- weighing and shaken to eliminate as much ferences in chemical content between the stipe possible of the surface water (Boariseth and and the frond.The atipe is fairly constant in 11aug, 1953), its chemical composition throughout the year, while -the frond undergoes marked seasonal vari- In Figure 14a the dry weight as a pareen- ations. tage of the fresh weiìt, or inversely the percentage water content, is shown for the frond Comprehensive studies on -the chemical colD- as it varies with season (Black, 1948a;Black position of L. hrperborea have been carried out and Dewar, 1949;Block, 1950a),There is by the Institute of Seaweed Research in Scotland clearly a maximum water content of about 87 per- and by the Noxieian Institute of Seaweed cent in May and a minimum of about 70 percent in Research,Fzrther references can be gained from Otober,This maximum occurs near the end of the series of publications from each institute fast growth when the frond has almost maxi mal and in Levriri,g et al. (1969). surface area and minimal storage material, while the minimum occurs at the end of the summer 7.1Water content assimilatory period when reserves are highest. BecauBo most chemical analyses of algae are As Been from Figure 14b, stipes are leen expressed in terms of percentage of dry weight, variable in water content, differing little the fresh weight has not often been determined from 85 percent (15 percent dry matter) throu at the same time so the dry weight as a percentage out -the year (Black, 1948a;1950a),Older of fresh weight, or inversely the water content, a-Lipes tend to have more dry matter than younçr hararely been reported.Because of adhering stipes (Black et a]., 1959).

30 70

Fie. 14Total dry matter expressed as a percentage of fresh weight, or water as a percentage of fresh weight (right hand scale), in L. hyperborea fronds (a) and stjpee (b).

Ash expressed as a percentage of fresh weight in fronds.Data replotted from: Black (1j48a) B1ak and Dewar (1949) Black ( 1950a) ncon and Ha91(19'ó) P2 FI ¶ /387 Laminaria hyperboroa 7.2Inorrçunio oontituentei mlninaue of 0.34 percent of total dry matter in fronde and 0.48 percent in stipes and a mean The total ash left after incineration and menimwn of 0.77 percent in fronds and 0.86 per- expressed as a percentaße of' the dry matter is cent in stipes.The levels were rather variahia shown for the frond in F'igure 15a and and. e-tipe and the minima and maxima not confined to autumn in Figure 15b (Black and Dewar, 1949;Black, and. spring. Larsen and 11aug (1961) found that 1950a;11aug and Jensen, 1954;Jensen end Hang, peripheral and older stipe tissue were higher in 1956).,Again there is considerable seasonal iodine.Typical con-tents of' acme inorganic variation in the frond (from about 15 percent constituents of spring and autumn samples of in autumn to about 35 percent in spring) 'Wi fronde and stipes are given in Table X (Jens mach lees in 'the atipe (32 to 40 percent)0 unpublished). This percentage variation is mainly due, however9 'to variations in the total dry matter and if Sorne do-terminations of' trace elements in the total ash of the frond is expressed as a L, h,'yperborea are shown in Table XI.,Those by eroontage of the fresh weight as in Figure 14a Black and Mitchell (1952) are means of three (Jensen and hug, 1956)9 i-t is shown 'to be measurements on material from fror.ds collected remarkably constant. in January and May,Those by Lnmde (1970) of fronds from Reine are means of six determinations Erly data on the composition of -the at two-monthly thiervais,The only elements ninerals in L. Ierborea tiere collected by showing regular variation with season were Vinogradov (l95e 11aug unii Jensen (1954) and arsenio, zinc and.ntIrnony, all lower from June Jensen and 11aug (1956) followed 'the seasonal to October than in 'the other montte,There seem chango in iodine concentration and in four to be considerable differences in the levels populations from different sites found a mean reported. by 'the two separate authors,

Fig. 15Ash expressed as a percentage of dry matter in L. hyperborea fronds (a) and stipes (b).Da-ta replotted from: Black and Dewar (1949); Black (1950a); and from 11aug and Jensen (1954) and Jensen and Haag (1956); ), Reine; ', Vard'c$; Jbpevaer. FII4 S8Laxninaria lierborea

TABLE X Inorganic cons-tituents of L. hperborea, in percentage of dry matter (Jensen, unpublished)

Frond

0.3 0,6 0,6 0,8 K 8.4 11.1 Na 3.2 1.6 Ca 1.4 1.7 Mg 0.8 0.6 Fe 0.03 0,028 0,018 0.01 504 4.6 2.7 PO4 0,6 0.5 Cl 8,8 11.7 Br 0.1 0.1 Cu 0,001 0.002 0,002 Zn 0,012 0,02 0,012

T/J3LE XI Trace elements found in L. rborea fronde in nkg of th'y matter

Black and Mitchell (1952): Lund.e (1970): Jensen (Tab.e x)

Black and Mitchell (1952): Thuido (1970): Jensen (Table X): F'I1 S8Laminaria erborea 7.30rjanic constituents Figure 16b and o.Here the variation in the fronds is even more marked, thou* the suiaser 7.31Carbohydrates rise cornea later than in mannitol.ThuR as a reserve it is laid down later -than mannitol ûnd The main carbohydrates of both fronds and probably depleted later.Black (1950e) found stipe of L. hyperborea are (in order of abundance) that during the period -that -the laminaran con- alginlo acid, laminaran, mannitol, cellulose and ton-t of the frond is high -there is a decrease ftzcoidan, in the amount of this substance with the depth of occurrence of the plant.A me-thod to isolato The simples-t, mannitol, is a fairly early glucose from L. hyperborea has been described photosynthetic product in Phaeophyceae and may byBlack e-t aL (1953, 1955), be the main respiratory substrate as has been shown -to be likely in Fucus (Percival and Fuooidan la oomposed of suiphatod McDowell, 1967, p. 22).It is a storage carbo- fuoose units.In the Phaeophyooae it may be hydrate.The seasonal variation in manriitol as assooiateà i.iith resistance to desiccation, percentage of dry matter in fronds and stipes being highly hygroscopio, and is low im of L. hyperborea from several sites in Norway i Laminaria, being less than four percent of the shown in Figure 16a and b (Haug and Jensen dry matter (Black, 1954a).Black found that 1954;Jensen and Hau.g, 1956).There is clearly it was higher in the frond 1han in the a-tipo a marked seasonal change in fronds with a sudden and in both showed a seasonal variation similar rise in response to the bes-t months for irra- -to that of mannitol,I-b may, therefore, be diance, and a slow decline in the winter when another but minor storage substance in L. respiration probably exceeds photosynthesis, The seasonal variation in the stipes, where hyperborea.The isolation and. pu.rifioe.tion nionnitol levels are lower, is distinct hirt much is described by Percival and Roas (1950) laco pronounced,Black (1948b) was the first -to observe these seasonal variations and -the I-t should be noted here that all the ator.- frond and stipe differences and his later resulte e compounds in this species show a similar., (Black and Dewar1949;Black, 1950a, 1955b, though not identical, variation with season in Table II, Fig. 2) are in agreement with those in the frond.Where the new and old fronds have Figure 16, though higher mannitol levels in been separated (Haug and Jensen, 1954, Pig. 7), fronds were recorded,Methods for the isola- mannitol and laminaren in the old frond decline tion of mannitol from dry or wet Laminaria aro steadily -to nearly zero between February and described in Black, Dewar and Woodward (1951), June while -the levels in the new frond remain steady and climb rapidly after Juno,Because The polysaccharide laminaran (previously storage materials are at relatively low levelo called laminarin) is a hot water-soluble, mainly in -the stipe and vary li-tilo with season1 it 1,3-j3 -linked. polyglucaxi.Its properties and appears that this part óf -the plant does not constitution have been investigated by several function as -the main s-borage organ.Both these authors (Connel et al. 1950;Black, Cornhill, observations fil well wi-bk LUning's (1970a) Dewar and Woodward,1951;Black and Dewar, 1954; observations that muoh of the early growth of ?riedlaender et al,1954; /inderson et al., the new frond is dependent on the presence of 1958;Minan et nl,, 1965a, 1965b;Fleming et the old frond but not of -the stipe (see 2.41). al., 1966).There are -two forms, the "insoluble" The old frond is thus the storage organ. which precipitates spontaneously in cold water and the "soluble" which does not because of more Aig-mio acid is a polytironide peculiar to branches in the chains (Fleming end Hanners, the Pbaeophyceae.It is composed of D-san- 1965).Laminaran from L. is predomi- nuronic and. L.-guluronic acid unite and 11aug nantly insoluble, while that from L.jitata (1964) showed that i-b in a mixture of hater-c- and. L. sacchnrina is mainly soluble,Peat et polymers of these, the proportions varying ai,, (1958) showed thai the insoluble fario con- between aig-mates from different sources, for tained about two percent of mannitol, these example -the new and old frond of L. hyperhorea units being at the ends of the chains and also (Hang et al., 1969) and between stipes from that there were some 1,6-í3-links between the different sitea.Ion exchange and. acidic glucose units.Further work on -the struciure of properties were shown to depend on -these pro- laminaran is eumxnarized by Percival and McDowell portions (Hang, 1964).Quite a lot la known of (1967, pp. 55-65).Its synthetic pathway has the synthetic pa-bhway of this eubs-tance and no-t been elucidated but there is a suggestion this is outlined by Percival and McDowell -that nanni-tol and laminaren are readily inter- (1967, pp., 17-19).I-t is likely that tho path- convertible (Percival and McDowell, 1967, p20). way differs from that of nenni-bol.Alginic It is a reservo material common in Pìaeophyoeae. acid mainly occurs in intercellular spaces but The seasonel variation in laminaran, in plants in also pronom-t as a major component on cell from the same source an analysed for mannitol, walls, being situated in the middle lame1l i-ni is shown for fronds and stipes respectively in the primary membrane, outeide 'the celulone (Anderson, 1956).Andersonhowsd that the FIitS8 Laminaria erborea

9 20 A A A MaJnitoI $ A 5 A q. t b ¡ A o o IO A A A t o f . Q A q. A A A t G t G A A G : * t:'

30 Laminaren C d * ¡A A

A A 20 A Q A G t s . $ A G

o o G $ G o A q. * q. q. q. . Q o o A O o 5 q. O q. 0 4 o O A A O A i * A t L * A u q. z q. o A 30 A o A 4 4 $ q. 4 q. A 4 A 1g i flic cid G $ : q. $ . q. s o A * q. q. a A o A A $ A ¡ t, 20 O G

e f I0

F M A M J J A S O N DJ F M .A M i A So N D Pig. 16Xannitol (a, b), lwninaran (o, d) and o]inio anid (o, f) oe& sa s peroent of dry mat-ter in L. hyperborec fronde (a, o, o) and $tipee (b, d, f).Data reploitod from Rang c.d. Jcnnen (1954) an(1 Jeneen and Haug (1956), Reine; V i; ,Repevaar. 7 6 FIR1. 387 Laminaria erborea proportion of al,inic acid increased toward the t3 to 15 percent around April, though ¿U percent centro of the tissue of Laminaria, the thick has been recorded in new fronds at that tirso walls of the medulla thus mainly consisting of (Hang and Jensen1954;Jensen and Hang, 1956), alginic acid rather than cellulose.Larson and This seasonal variation (also reported by Black Flaug (1961) also found an increase in alginate and Dewar, 1949 and Black, 195Oa) in similar to toward the centre of the stipe. that of alginic acid and presumably also reflects the variation in storage polysaccharidoe making The levels of aiginic acid found at dif- up the bulk of the dry weight in late summer, feront seasons in fronds and stipes in Norway Thus, when expressed as a percentage of fresh from the same sites as analysed for mannitol and weight, frond protein varied from one percent in laminaran are shown in Figure 16e and f.There June to two percent in December (Jensen and is clearly a marked seasonal variation in the 11aug, 1956), a cycle similar -to that of alginio frond, varying from around 20 percent of the dry acid again,The protein content of the stipes matter in autumn to 30 percent in spring.The varies little with season aM umully lies stipea, however, show no seasonal variation but between 7 and 12 percent of the dry matter a higher alginate content of about 30 to 37 per- (Black, 1950a;Haag and Jénsen, 1954;Jensen cent of dry matter.Black and Dewar (1949) and and Haag, 1956). Black (1950a) observed much lower values for alginic acid of 8 to 10 percent of dry matter Although most of the nitrogenous matter in in the fronds and 19 to 23 percent in the stipes. L. ¿erborea is in the form of proteins and. The seasonal variation in alginic acid shown by peptides, some freecc-.amino acids have been the fronda can be considered as a reflection of found, mainly alanine, aspart±o and glutamic the seasonai variation in the storage compounds acids (channing and. Young, 1953).The amino-acid. rranitol ed land.riaraxi because all are expressed composition of isolated bulk proteins from L. as a percntage of dry weight.When frond hyperborea fronds were analysed by Coulson t1955). alginiá acid is expressed as a percentage of CoulsonIT953) observed a seasonal variation in fresh weih-t (Jensen and Haug, 1956, Fig. 13) cys-teic acid content fi-orn nil in September to a this pattm disappears and alginic acid shows maxi mum between Hay and August. ¡a jnanjmujn in December and a minimum in Juno, which may or may not be significant.As there 7.33Fat is little variation in storage compounds in the stipe and their levels aro low, alginic acid The ether-ertraotable matter (total fat) is varies little with season and its percentage is low in all Laminaria species.In tho fronds of correspondingly high.The viscosity of sodium L. prborea it varies from about 0.5 tô 1,5 alginate from frond and stipe respectively and percent of the dry matter, usually being lown after different treatments was invostigated by August and high in the springwhile in the stipes Black et al, (1952). it differs little from 0.5 percent (Haag and Jensen, 1954;Jensen and Haug, 1956). Cellulose, consisting of 1,4-ß-linked glucose units and, similar to the higher plant 7.34Pigments substance (Percival and Ross, 1949) is present Th5 Phaeophyoeae contain ohlorephylls a in the cell walls of L. hyperborea in much smaller and o, fuooxanthin, violaxanthin andj3-carotene quantities than alginic acid.Black (i950b) (Jensen, 1966).Ssybold and Egio (1938) observed the seasonal variation in the frond analysed L, orborea and found 2 320 ing of and stipe.It varied in the frond from about chlorophyll, 85 mg/3 -carotene, 469 ing Ñcoxan- four percent of the dry matter to nearly six thin and 45 mg "xanthophyll" per kilogramme of percent.In the atipe the levels varied from dry matter.owen (1954) made detersinationa of eight to ten percent of dry matter. frond and stipe materiaL, fresh or previously dried, but all his fiures aro considerably Ehzyines capable of decompoeing insoluble lower than those of Sybold and Egle Ris laminaran are prodncecl by many micro-organisms values for L. digitata ai-e similarly much lower such as those of the microflora of the bovine than those of both Seybold and Egle and of P.lmen. The decomposition of alginates is not so Jensen (1966), who did not make determinations widespread a phenomenon,Laminarais and an on L. byperborea.Jensen found a level for alginate have been isolated (Chos-ters ei , violaxanthin in Le diitaia of 110 ms/kg dry 1956). matter, 7.32 Proteins 7,35Other organic constituents The protein content of brown algae is generally quite low and that of L,rperborea A number of vitamins have been shown to is no exception.There is a seasonal variation occur tuL. hyperborea.Vitamin A which in iblU in the protein content of fronde from about case only occurs in the form of ita pro- five percent of the dry weight around August to vitamin ß-carotene, is dealt with in the previous eec-t ion, Of the vttamnìfl B group I/8 Laminaria arborea

the niacin content of this species was invea-- borea which should correspond to about 700 to tigated by Larsen (1958) over a period of six 3 400 mg/kg dry matter,The ni.ximum wan in years.There was a remarkably consistent seasonal .Aril.R3nnerstrand (1943) found 70 mg/kg wet variation in the fronda from about IO mg/kg dry wei..-t in summers oorresponcUng to about 350 mgI matter in October -to about 35 mg/kg in April and kg dry weight.Of -the vitamin E group only a single year's obsera-tions on the levels in the .0-tocopherol has been found, in small but con-. stipe indicated a srnilar cycle.A similar but ale-tent quantities of 10 bo 30 mg/kg of dry much less regular seasonal variation was observed matter (Brown, 1953;Jensen, 1969). in the bio-tin content of the fronds, varying from about 0.3 to 0.5 mJk of dry matter (Larsen, Fucos-berol, the typical steroid of brown 1961).KarlatrThn (1963) measured the iritaniin B12 ale, occurs in L. erborea as approximately content of L. hyperborca and found 0.04 mg/kg of 0,1 percent of the dry matter (Black and Corn- dry matter, bwt Ericeon (1953) found up to i mg! hill 1951). kg in L. maccharina,Erioson also found vitamins B1, B2, folio and folinic acid in. uxins bave been detected in this species, charma,Lwide and Lie (1938) found vitanin C in chromatogram spots close to bu-t possibly not levels of 100-470 mg/kg wet weight of L. hyper- identical with indolylace-bic acid cnd indoly- lacetoni-brile (Nowa-b, 1964), FIF(M/S8Laminaria erborea 8:1 8 UTILIZATION 8.3 Nanure 8.1Human food Seaweed has been used as manure by ooasta]. farmers for centuries.L. hyerboraa is parti- This species is used very little as human cularly useful in some areas because large quan- food.In the past, in times of starvation, the tities are washed ashore at times.It is still frond has been eaten in Iceland (Haussen, 1964). used occasIonally In this way in Scotland and other places though generally other forme of 8.2Animal fodder fertilizer have taken its place. There IB similarly little use for L. Seaweed meal or liquid fertilizers have a h.yperba as animal food though cattle occa- great many applications and advaxttous side sionally eat cast plante on the shore,, effects (Booth, 1966) but again these are preparedmainly (in1ropo) from Aacophyllwn and There have been many experiments on the use not L.,yperborea.Sôme liquid fertilizer is of dried seaweed meal as a supplement to the manufactured from L. hyperborea fronds in diet of domestic animals and although most of Britain, however.An example of an experiment the meal used for this is prepared from involving the use of this species as fertilizer Itaco,phyllum nodostzm (see. Baardseth, 1970) Borne is that by Myklestad. (1964).Ii was found that, experiments have included L. hyperborea meal. as a supplement to basic fertilizing of newly For example, Hand (1953),Black (1954b) axt cultivated ground, seaweed stimulated an increase Hand and Tyler (1955) found that replacing In turnip production which was greater than ten percentf the normal diet of egg-laying could be attributed to the provision of' N, K and hens with L. i,erborea meal (preferablyxe- P,On solle in good condition, however, it pared from Irondshad, no harmful effect (though little effect, water consunption went up), but did not appear to have an advantage in an alrea1y ba].anoed 8.4 Industrial products diet.On the other hand, Hphe and Sandvi]c (1956) chowed that L. hyperborca meal added to The carlies-t industrial application for a diet noi supplemented with yeast, grass meal this species was in -the production ofkelp", or cod liver oil stimulated a marked improve- This originaily meant the ash from burnt sea'Tood. ment in the growth and health of chicks. This izas used as e. source of soda, particularly Particularly good resulte were obtained from for the class iadustry (C'aapman, 1970).' Soon meal prepared from young fronds,However, en after altornr.tivo sources ucre found for this, addition of five percent L. ,erborea meal to crly in the rinetoon h oentn'y, the ach wan an all-round basal diet for chicks, alhouh ucd íuctoado a coures of iodine and this con- doing no harm, had no beneficial effect (Hie and tinuecior ahot a een'ury in Scotland,In Sannan, 1958).Ten or 15 percent additions wore Frar ehoiovov, ii; continued for longer (Feld-' in fact unfavoux'able.Similarly the addition of win, 1953) bri ceased In 1958 (Ohapmu, 1910). ten percent L. hyperborea meal to the dl.et of dairy cows had no effect on milk production Nost.of the L. porbopoc noi' collected for (Burt et al., 1954).Black and Joodward (1951), OOTtnercial purposes in No)C and Brjain je from theoretical considerations, concluded uoed for the manufacture of alginaton.Thin that seaweed mea]. (including thai. from L. species is 3itle used. b' alginate industries in ), supplied as ten percent of the diet, other coìintrioa,In fl'ítin about one third of could provide enou:' NaCl and 12in addition -the algina-te produoed originates from L hyper- to Cu, Co, Mn, Fe and other possible trace borec,The Br&1.nh &ncìnslzy (Îtiginaie Indus- elements and vitamina for the normal requtromeute tries Ltd., London) usos c-ir'dried stipes while of all farm livestock. -the PotrogIcm f'nstory (Ptan &Fagortun A/S, Prammen) utii&;oc thehole plant, undriod.The Swule (1956) found that ihon leyinbons e.ginl,c acid is ettracted from the algal material were given seven percent of their feed as L. with h'ise, precipitated as free acid and neu-trz- hyperborea meal the Icontent of the eggs ias llaed -to sodium alginate ithioh is the main 150 times that of those fed on the usual diet. product.Aid-miO acid and ita salts, the algi-

sates, find very many applications in pharms-- Although the use of seaweed meal in livestock ceutical and food industries and in printing and feedIng has many advantages (Baardscth, 1970), paper sizing processes (see Chapman, 1970). that frein L.yoorborea hAul too high an iodine content to be of general uso end is not sold for There has been quite a lot of work on the this purpse. use of the polysaocharido laminaran in medicine. The insoluble forzi can be nest economically Preservation by ensuing has been studied isolated from L.rperborea fronds between (Black 1955a). Augus-t arid December (Reid, 1956).Ii cari act as 8:2 FII* SBLaminaria hperborea an e icoauThrr after beingsulphatecl (DeuarD Mamo et al., 1958).Aflhough laminaran han 1956).With about 0.6 sulphate g:roupe per' advantages as alown powder substitute, the glucose molecule this lsminarn can act as an insoluble form(economically producedfrom L. antilipaemio tnj (Betorman and Eve.ne 1957) hyperboreaonly) was foundtoaffec'liver'and Horcver, somelaincrtn sulphaec have bean found kidney tissue of test animals and its :use wan io have harmful effoste (Mamo end. Thorpe, 1957; abandoned(B],aine,1951) FIRM/5487 L.vniimria hyperhorct 2!J.. 9 rurERErrc'.1s

Athains, S.S. and H.N. Thorpe,The anti-coatilant activity and toxicity of laminarin sulphate. 1957 J.Pharm.Lond., 9:459-63 Adams, S.S., H.M. Thorpe and L.E. Glynn,Effect of n iaminax'in eui.phate (LM46) on bone growth. 1958 Lancet, Sept. 20:618 Anderson, F.B. et al.,The constitution of laminaria,Part 3.The fine strtioturo of insoluble 1958 laminarin.J.Chem.Soo., 1958 (663):3233-43 Anderson, G.,On the detection of alginic acid in tissues by means of birefri:ngnnoe.Proc.Int, 1956 Seaweed Symp., 2:1 19-24 Annan, W.D., E. Hirsi and D.J. Manners,The constitution of laminaria.Part 4.The minor component 1965a sugar.3.Chern.Soo., (34):220.-6 The constitution of laminarin.Part 5.The location of 1,6-glucosidic linkages. 1965b J.Chem.Soc., 1965:885

Arcnchourg, J.E., Observationes Phycologioae,4. No Acta R.Soo,Sci,TJ:s. 12 1883 Baardoeth, !.,Kvn.ntitative -tare--underskalser i Lofoten og Salten sommeren 1952.Rep.Norw.Inst. 1954 Seaweel Res,, 6:47 p. A method of estimating the physode content in brown ale, Rep.Norw.Inpt.Seaweed Ros., i98 20:1-6 Anatomical investigations of some brown algae.Proc.Inb.Seaweed Symp,, 4:39-41 164 Synopsis of biological data on knobbed wrack Ascophylluzn nodosuin (rAnnaeua) Leolis, 1970 FAO Fish,Synops., (38)Rev.j:pag.var. Baardseth, E. and A. Hang,Individual variation of sorno constituents in brown algae, reliability 1953 of anaiytical results,Re ,Norw.Inst.Seaweed l'es., (2):23 p. Bellamy, DJ. e-b al,,Effecta of pollution from the Torrey Canyon on lit-bora]. and sublittoral ecotaye- 1967 tema,Nature, Load,, 216:1170-3 The place of ecological monitoring in the study of pollution of +,he marino env'ironn'ont. 1970 Paper presented to FAO Technical Conference on Marine Pollution and its Effects on Living Resources and Fishing, Rome, 9-18 December 1970, FIR:MP/70/E-65:12 p, Bellamy, D.J., D.M. John and A. dhittiok,The "lelp forest e000yi ìs a "phybometer" in the study 1968 of pollution of' the inshore environment, 1968:79-82 Beutei'rnan, E.M.M. and J. Evans,Anti-lipaemio agent without abi-.00agulant action.Bz'.med,J., 1951 50141310

Black, W.AP.,Seasonal variation in chemical oonmti'wtion of como of the rsablittorei seaweeds o on 1948a to Scotland.Part 1, Laminaria olouctoni0J.Soo,Ohom.a.d, 67:165..-8 Seasonal variation in chemical eonstitution of some Commonit: Nature, tond., 161:174 The seasonal variation in wei(!ftit chemical ootnpoeitioa of the o ori Bri-tieb 1950a Lwninartaoeae.JMar0Bio].Aeaoc,U,X,, 29(1):45-72 The ecasonal vz'intion in the cellulose content of the common Scotti inariaceso 1950b and 1'ueaooae,joMrir.Boj,AranooeU,J(., 29(2):379-87 The effect of depth of' immernion on the chemical constitution of nome of the' sublittoral 1950e seaweeds common to Scotland.J. oc.Chan,Xnal., 69,161-5 2:2 FI,Rl/S87 Laminaria hyperborca Black, W.A.P,The seasonal variation in the combined L-fucoae content of the corrsion British 1954a Laxrrinariaceao and Fucaceae,J.Sci.Food Agrio., 5:445-8 Seaweed as poultry food.Wór].d Poultry Sci.J., 10:33-5 1954b The preservation of seaweed by ensilinand bac-tericides.J.Soi.Food Agrio., 6:14-23 1955a Seaweeds and their constituents in foods for man and animal0Chem,Ind., 1955:1640-5 1955b Black, W.A.P. and 1,J. Cornhill,A method for the estimatión of fucosterol in seaweeds.J,Sci.Food 1951 Agrio., 2:387-90 Black, W.A.P. and LT. Dewar,Correlation of sorné of the physics]. and. ohemioa]. properties of the ses 1949 with the chemical constitution of the algae.J.Ma,r.Biol.Assoo,U,K,, 28:673-705 The properties of algal chemicals0 2.Some darivates of laminaria.J,Sci.Food Agrio., 1954 5:176-83 Black, W.A.P, and R.L, Mitchell,Trace elements in the common brown algae and in seavater,J,Mar, 1952 Biol.Assoo,U.K., 30:575-84 Black, W.A.1'. and F,N, Woociward.,The value of seaweeds in animal feedinstuffs as a source of 1957 minerals, trace elements and vitamins.Emp.J.Exp.Agrie., 25(97):51-9 Black, W,A.I'., W.J. Cornhill and. LT. Dewar,The properties of the al chemicals,1.The evnlu.a?- 1952 -tion of -the common British brown marine algae as's source of alginate.J.Sci.Food. Aric,, 3:542-50 Black, W.A,P,, E.T. Dewar and F.N. Woodward,Manufacture of algal chemicals. 2.Laboratory-scale 1951 isolation of mannitol from brown marine algae., J.Appl.Chem., 1:414-24 Manufacture of alga]. chemicals.5.Laboratory-scale isolation of D-glucose from 1953 laminaria. J.Sci.Food Agrio., 4:58-64 Manufacture of algal chemicals,7.The isolation of algal carbohydrates by meanscf 1955 charcoal columns,J.Sci.Food Agrio., 1955:74-63 Black, W.A.P, W.D. Richardson and F.T. Walker,Chemical and growth gradients of Laminaria clourr toni. 1959 Edm, (L.yperborea Foal.).Econ,Proo.R.Dublin Soc, 4(8)1137-49 Black, W.A.P. et al.,Manufacture of algal chemicals,3.Laboratory-scale isolation of laminaria 1951 from brown marine algae.J.Appl.Chem., 1:505-17 Blame, CL,Laminaria as a talcum powter substituto.Med.Press, 226:611-2 1951 Boney, A.D.,Aspects of the biolor of the seaweeds of economic importance.Adv.Ma.r.Biol,, 3:105-253 1965 Booth, E.,A method of drying seaweed using a steam-heated drum dryer.J.Soi.Food Agrio., 7(11) 1956 705-10 Some properties of seaweed manures.Proc,In-t,Seaweed. Sy-mp., 5:349-57 1966 Breivik, K.,Observations on the macroscopic algal vegetation in the fjords near Siavanger, Norway, 1958 Nytt Ma,Bot., 6:19-37 Drown, F.The occurrence of S-t000pherol in ceaweed,Chem.Ind., 1953:174 1953 FImt/38' Laminaria herborea Bruce, J.R,, J.S. Coleman and N,S. Jones (Eda),Marine fauna of the Isle of Man and its surrounding 1963 seas.Liverpool, University Press, 307 p.

Burrows, E.M., Anexperimental assessment of some of the characters used for specific delimitation in 1964 the genus Lantharia.J.Mar.Biol.Aesoo.U.K., 44:137-43 Synopsis of biological data on Laminaria sancharina.FAO Fiith,Synops., (in preparation)

Burt, A.W.A., S. Bartlett and. S.J. Rowland, The use of seaweed meals in concentrate mixtures for 1954 dairy cows.J.Dairy Res., 21:299-304 (manning, D.M. and G.T. Young,Amino acids and peptides.Part 10.The nitrogenous constituents of 1953 some marine algae.J,Chem.Soc., 1953(503):2481-91 Chanan, V.3.,Methods oi' surveying Laminaria beds.J,Mar.Biol.Aesoc.U.K., 26:37-60 1944 Seaweed resources along the shores of Groat Britain.Econ.Boi., 2:363-78 1948 Seaweeds and their usos.London, Mothuen, 304 p 1970 Cheaters, C,G000, A. Apinis and M. Turner,Stùdies of thé decomposition of seaweeds and seaweed pro- 1956 ducts by micro-organisms.Proc.Linn,Soc.Lond., 1956:87-97 Connell, 3.1., E.L, Hirat and. E.G,V. Percival,The constitution of laminaria,Part 1.An investi- 1950 gation on laminarin isolated from Laminaria cloustoni,J.Chern.Soc., 1950(688):3494-3500 Coulson, C.B.,Amino-acids of marine algae.Chem.Ind., 1953:971-2 1953 Plant proteins,i.Proteins and amino-acids of marine algae.J.Sci.Food Agrio., 1955 6:674-82 Davis, F.M.,An account of the fishing gear of England and Wales.Fiah,Invost., Lond.(2), 21(8):1ap 1958 Do Toni, J.B.,Sylloge algarum omnium huousque cognitarum.Padua 1889-1924 Dewar, E.T.,Sodium laininarin as a blood coagulant. Froc,Inj,Seaweed Symp,, 2:55-61 1956 Dizerbo, A.H.,Algues du Spitsberg rcoltes par l'Expdition geographique Arinick Moign (1964). 1969 Rev.Algol., 9:297-8 Drach, P.,Premiares recherches en scaphandre autonome sur leformations do laminaires en zone 1949 littorale profonde.C.R,Somin.Sanoe Soc.Biogogr., 227:46-9

Drtzehl, L.D.,Taxonomyandistribution of northeast Pacific species of Laminaria.Can.J,Bot., 46 1968 539-47 Du Rietz, G.E., Studier 5ver de skandinaviaka Laminaria-arterna.Bot.,Not., 1920 1920 Nomenclature of Laminaria digitata (Ruda.) Laznour, Laminaria hyperborea (Chum.) 1953 Foalie Sveriek Bot.Tidekr., 47:1 Ecbnondston, T.,A flora of Shetland, comprehending a list of the Shetland Islands, with remarks on 1845 their topography, geology and climate etc.Aberdeen Ernst, J.,Sur la vgtation sous-marine de la Manche d'aps des observations en scaphandre autonome, 1955 C.R,Hebd,S&inces Acad,Soi,, Paris, 241:1066-68 FIRM s87 Laminari J,erbo Ericaon, L.E..,On the Vitamin B12 -, Folio Acid, and. Follaje Acid groups of factors, and on the occu.r- 1953 rence of these vitamins, and of Niacin, Pantothenic acid, and amino acids in a number of marine algae.Uppsala, Almquist and Wikeells Boktrykkeri AB Evuna, L.V,,Cy-tological stwlioe in the Laminariales,Ann.Bob,, 29:541-62 1965 Feldmann, J., L'utilisation dea algues marines on France,Proo.Inb.Seaweed Symp., 1:88-9 1953 Feldman, J. and R. Lazni,Flore e-b vg-bation marines de la oste basque franaiae. Bull.Soc.Bot.Fr,, 194L 88:123-42 Fleming, M. and P.3. Manners,A structural difference between, the soluble and. insoluble foz,ns o' 1965 lazainarin,. Bioohom.J., 94:17 p. Flood, D.T.,Seaweed resources of Eire,Proo.Int.Seaweed Symp., 1:87-8 1953 Forster, G,R,,Underwater observations on rooks off Stoke. Point and Dartmouth.J.Mar.Biol.Assoc,U.X., 1955 34:197-9 Foche, M.,Bidrag til kundekaben ont de -tugruppen Digitatae hrende Lazninarier.Forh.Videaskaselsk. 1884 ICristiania, 2:19-32 lus Ueber die Laminar-Len Norwegens.Forh.Videnakapsselsk.Xria-biania, 1884(14): 1-112 Friedlaender, N.rL.G,, W.H, Cook and W.G. Mar-bin,Molecular weight and hydrodynamic properties of 1954 laminaria.Biocbim.Biophys.Aoba, 14: 136-.14 Gardner, R.G. and 'P.3. Mitchell,Throuis-ciroulation drying of seaweed,1.Laminaria clouabon3. 1953a a-tipe,3.Soi,Food. A,o., 4:113-29 Through-circulation drying of seaweed. 2.Lantin9 'ìa cloustoni frond,J,Zci .Food Agrio,, 1953b 4:237-45 A study of seaweed drying.Proo.Int.Seawe t Symp., 2:63-81 1956 Graham, A. and V, Fretter,The life history of Patina pelluoica (L).J.Mar.Biol.Assoo,U.K., 26: 1947 590-601 Grenager, B,,Kvantibativu underspfkelser av taceforekomcter p9 Kvitsy og Karwy 1952.Ro2,Norw.Inst, 1953 Seaweed. Res., 3:53 p.

Kvnntitative undorsjkelser av tareforekomaterp9Tustna 1952 og 1953.Rep,Norw.Inat, 1954 Seaweed Res., 5:33 p. K-vantitative undersjkelser av tareforekomater i sr-Helgelanil 1952 og 1953.Rep..T4orw. 1955 Inat.Seaweed Res., 7:70 p. Kvantita-tive underskelser av tareforekomster i Øst-Finmark 1953.Rep.Norw.Inst,Soawoed 1956 Res., 13:37 p. Mvanbita-tive unders&elser av bang-. og tareforekomater i }Iely, Troms 1953.Rep.Norw. i953 Inst.Seaweed. Res., 21:1-31 ICvantitative unclersjlkelser av tang- cg taroforokoinstex' i Nord.-Frya he'rett 1954 og 1964 J,*ssund herred. 1956.Rep.Norw.In:t.Seaweed Res,, 28:53 p. FIRN/S87 Laminaria hyperborea Guignard, L.,Observations sur l'appareil mucifre des LaninariacosAnnScioNat.(Bota), 15:1-46 1892 Gunnerus, J,E,,Flora Norvegica.Nidrosiao 1766 Halleson, 3.V,,The uses of seaweeds in Iceland.Proo.Int.Seaweed S, 4:398-405 1964 Hand, C.J.E,,Feeding seaweed meal to laying hone.Proo.In-t.Seaweed Symp,, 1:68-9 1953 Hand, C.J,E. and C. Tyler,The effect of feeding different seaweed meals on the mineral and nitrogen 1955 metabolism of laying hens.J.Sci.Food Agrie., 6:743-54 Harnee, R.,An inve-bigation by cultural methods of some of the factors influencing the develornient 1932 of the gaxnetophy-tes and early etage,s of the sporophy-tes of Laminaria digitaba, L. saacharina and L. cloustoni.,Ann,Bo-b.,, 46:893

Hartog, C. den,The opilithic algal communities occurring on the coast of the Netherlands,,Amsterdam, 1959 North-Holland Publishing Co,, 241 p. Harvey, L.A.,The marine fauna and flora of the Isles of Scilly.The islands and their ecology. 1969 J,Nat,His-t., 3:3-18 Harvey, W,H.,Phycologia Britannica.London, Reeve & Benham 1846-51 Hasagawa, T.,Cultivation of Laminaria in Japan.Proc,Pao.Soi.Congr,, 11 (7): 19 1966 11aug, A.,Composition arid properties of alginates.Re ,Norw,Inst,Seaweed Res,, 30:123 p, 1964 Hang, A. arid A. Jensen,Seasonal variations in the chemical composition of Alaria esculenti., 1954 Laminaria sacohanina, Laminaria byporborea and Laminaria digitata from northern Norway. Rep.Norw.Inst.Seaweed Res,, 4:14 p. Hang, A., B, Larsen and E. Baardseth,Comparison of t10 constitution of alginates from different 1969 sources.Proc.Int,Seaweed Symp., 6:443-51 Hay, J.L,Harvesting of brown sublittoraleaweeda,2,The continuons puol method,Engineer,, 1952 Lond., 193:814-7, 846-51 Holland-Hansen, B. and F. Naneen,The sea west of Spitzbergen,Skr,Vidensk,,ChxistianiaMat,-Nat, 1912 1912(12):89 p. Reek, C. van den and M. Donze,The algal vegetation of the rocky C8te Basque (sw France).Bull,Cont, 1966 E-tud,Rech.Sci,, Biarritz, 6(2):289-319 114e, 3. and Ø. Sandyik,Experiments with seaweed meals as supplements to rations for chicks and 1956 laying hens,Proc.,Int,Seaweed Symp., 2:82-97 H4e, J, and F. Sannan,Further experiments with seaweed meals as supplements to rations for chicks 1958 and laying hens,Abs-tr.Int.Seaweed Symp,, 3:48-9 Horkin, R. and J.M. 1ain,A preliminary report on the effect of marine pollutants on Laminaria 1971 hyperborea.Mar.Pollut,Bull., 2:75-7 Jaasund, E,,Aspects of the marine algal vegetation of north Norway.Bot.Gothoburg, 4:174 p. 1965 Jackson, P.,Director's report, engineering division. p,Scott,Seaweed Res,kssoo., 1948:23-30 1949 9:6 FIRM/S87 Laminaria hyperborea Jackson, P.,Harvesting of brown sublittoral seaweeds.1.Assessment of the problem and review of 1952 preliminary aperimentm.Engineer, Lond,, 193:716-20, 750-51 Harvesting machinery for brown sublittoral seaweeds.Engineer, Lond., 203:400-1, 439-41 1957 Jackson, P. and I.B. r4acduff,Harvesting of brown sublittorai seaweeds.4.Entrainment of plants. 1952 Engineer, Loud., 194:42-7 Jackson, P, and. R.F. Molver,Harvesting of brown sublittoral seaweeds.3.Use of reoiprocating cut- 1952 term. gineor, Load,, 194:2-5 Jackson, P. and R. Wolff,Sublittoral seaweed harvester.Research, Lond., 8:435 1955 Jensen, A.,Carotenoids of Norwegian brown seaweeds and of seaweed meals.Rep.Norw.Inst.Seaweed Res., 1966 31:138 Tocopherol content of seaweed and Beaweed meal,1.Analytical methods and distribution 1969 of tocopherols in benthic algae.J.SciJood Agrio., 2O:449-53 Jensen, A. aid A. Haag,Fargereakmjon tuadskillelE;e aStortare (Laminaria cloustoni) og Fingertare 1952 (Laminaria digitata).Tidsskr.Kjemi Bervves., (8):138-9 Geographical and seasonal variation in the chemical composition of Laminariajierborea 1956 and Laminaria digitata from the Norwegian coast, 20Norw.Inst.Seaweed Res., 14:; p. Jobn, D.M.,Studies on littoral and sublittoral ecosystems.Thesis, Durham University,168 p. 1968 An ecological study on Laminaria ochx'oleuoa.J.Mar.Biol.Assoc,U,K,, 49(1):175-87 1969 Johnston1, C.S, and J.M. Davies,Aspects of the intermediary metabolism of members of the Fu.calea and 1969 Leininarìales,Proc.Int,Seaweecl Sym, 6:501-6 Jones, ff.S. and J0M, Kam,Subtidal algal colonization flowing the removal of Echinus.Helgolaonder 1967 tiaa.Meeresunters., 15:460-6 J6nsson, H., The marine algal vetation of Iceland.In The Botany of Iceland by L.K. Rosenvinge .4 1912 E. Warming,London, Wheldon, Vol. 1:1-186 Jorde, I.,Algal associations of a coaetal area south of Bergen, Norway.Sars:.a, (23):52 p. 1966 Kam, J.M.,Direct observations on some Nansublittoral algae.J,Mar,Biol,Assoc.U.K., 39(3),609-30 1960 Algal speoiss lists:some sublittoral records.Br.Phycol.Bull,2:80-6 1961 Aspects of the biology of Laminaria hyperborea.i.Vertical distribution.J,Mar.Biol. 192 Assoo.U.K., 42(2):377-85 Aspects of the biology of Laminaria hyperborea.2.Age, weight and length.J.Mar.Biol. 1963 ÀSSOCOU.KO, 43(1):129-51 Aspects of the biology of Laminaria hyperborea.3,Survivul and. growth of gunetophytea. 1964 J.Mar.Biol,Assoc.U,K., 44x415-33 -_-- ,Aspects of the biology of Laminaria hyporborea.4.Growth of early sporophytes.J. 1965 }ar.Biol,Aesoo.U.K., 45:129-43 FIPJA/387 Laminaria hyperborea

Kam, J.M., The role of light in the ecology Of Laminaria hyperboroa. In Light as an ecological tac- 1966 tor, edited by R. Bainbridge, G.C. Evans and O. Raekham,Oxford, Blackwoll, pp.319-34

Populations of Laminaria hyperborea at various latitudes, Helgolnender Wise. 1967 Meereeunters.* 15:489-99

, The biology of Laminaria hyporborea. 5.Comparison with early stages of competitors. 1969 J.Mar.Biol.Asooc.1J.K., 49(2):455-73

The biology of Laminaria hy-perborea. 6. Some Norwegian populations. J.Mar,Biol. 197m Assoo.U.K., 51:387-408

Continuous recording of underwater light in relation to Laminaria distribution.In 1971b Proceedings of the 4th European Marino Biology Symposium, edited by D.J. Crisp. Cambridge, University Press, pp.335-46

Kam, .7.M, and P. Svendsen,A note on the behaviour of Patina pellucida in Britain and Norway. 1969 Sarsia, (38):25-30

Kanda, T,, On the gametophy-tes of some Japanese species of Laminariales,Sci0Pa.Ine-t.Algol.Ree. 1936 Hokkaido Univ., 1:221-60

-, On the gametophy-tes of some Japanese species of LaziinariaJ.es 5. Sci.Pap.InBt.Algpi. 1944 Res,Hokkaido Univ,, 3(1):121-54

Karletrm, O., The vitamin B12 content of maxine ialgae. Re ,Norw.Inst,Seaweed Res., 29 1963

Killian, K.., Beitr'ege zur Kenntnis der Laminarien, Z,Bot.,, 3:433-94 1911

Ki-tohing, J.A., Studies in sublittoral eoologye 2. Recolonization at the upper margin of the sublib- 1937 tora], region with a note on the denudation of Laminaria forests by storms, J,Ecol.., 25: 482-95

Studies in sublittoral ecology, 3. Laminaria forest on the west ooast of Scotland; 1941 study of zonation in relation -to wave action and illumination,Biol..Bull,Mar,Biol.[,ab.1 Woods Hole, 80:324-37

Knight-Jones, E.W., 11.0. Jònes and D. Lucas, A survey of a submarine rocky channel, Rep,Challeriger 1957 Soc,, 3(9):20-.1

Kuckuck, P., Boitrge zur Kenntnis der Moereoa],gen. 2, Zur Fortpflanzung der Phaeosporeen.Wise0 1912 Neeresunters.(AbtoHelgoland), 5:155-88

Kutaing, F., Pkyoologia generalis,Leipzig, 1843

ICylin, H., Studien fber die Algenflora der schwedischen Westkste,Upsala, Akadem. Abhanñlung 1907

La Pylale, A.J.M.B. de, uelques observations sur les productions do l'110 de Terre Neuve et our 1824 quelques algues do la cßte de France, appartenant au genro Laminaire.Ann.ScioNat.(1), 14

Larsen, B., The influence of season, habitat and age of tissue on the niacin content of some brown 1958 algae.. Rep.Norv,Inst.Seaweed Roe., 19:13 p,

The 'biotin content of marine algae.Re Norw.Inst,Seaweed Ros, 26:18 p, 1961

Larsen, B. and A. 11aug, The distribution of iodine and other constituents in stipe of Laminaria hypex- 1961 borea (Unna.) Foelie, Bot.lar., 2:250-4 2:8 FIRM/S87 Laminaria hyperborel Leighton, D01,., L.G, Jones and I.J. North,Ecological relationships between giant kelp and sea 1965 urchins in southern California.Proc.Int.Seaweed. Symy., 5:141-53 Le Jolis, A.,Examen des eopces confondues sous le nom do Laminaria digitata auct., suivi de quelques 1855 observations stir le genre Laminaria.M4m.Soc.imp.Sci.natChorbour, 3:241-312 Levring, P.,Reduced foxias in the Swedish marine flora.Proc,Int.Seaweed Symp., 1:20-2 1953 Loving, P., TL.AHoppe and 0.3. Schmid,Marine algae,A survey of research and utilization.Bot. 1969 .rrandb0, 1:421 p. L«aing, K.,Standing crop and leafarea index of the sublittoral Laminaria Bpooies near Helgoland0 1969a Nar0lJiol,, 3(3):282-6 Growth of amputatedand dark-exposed individuals of the brown alga Laminaria hyperborea, 1969b Mar.Biol,, 2:218-23 Cultivation of Laminaria hyperborea in situ and in continuous darlcness under laboratory 1970a oondtiions.Heßolaendar Wiss.Meeresu.nters., 20:79-88 Tauohimtersuohungen zur Vertikalverteilung der sublitoralen HelgolHnder Algenve:tation. 1970b Holgolaender Wiss.Meeresunters., 21:271-91 Seasonal owth of Laminaria rboroa under recorded underwater light conditions near 1971 Helgoland.In Proceedings of the 4th European Marine Biolor Symposium, edited by D.J. Crisp,Cambridge, University Press, pp.347-61

LuMe, G.,Analysis of trace elements in seaweed.J.Soi.Food 21:476-8 1970 LuMe, G. and J. Lie,Vitin C in Meeresalgen,Hoppe-Seyler's Z.Physiol.Chem., 254:227-40 1938 Lyngbye, B.C.,Tentomen Hydrophytologiae Danicae,Hafniae 1819 McAllister, H.A., T.A, Norton and E. Conway.A preliminaxr list of sublittoral marine al 1967 from the west of Scotland. ol,Bull,, 3(2):175-84

Marshall, W.,An underwater study of the epiphy-bes of Laminaria hyperborea (Gunn.) Foal,Br.Phyuol, 1960 Bull., 2:18-9 - Miyabe, K,,Laminarmaceae of Hokkaido0(In Japanese).In Nippon Kai s8-shi, by K. Okamura, pp.283-300 1936 - On the Laminariacoae of Holckaido,J.Sapporo Agrio.Coll., 1:1-50(in Japanese 1902) 1957 Miyabe,K, and M, Nagai,Leminariaceae of the Kurile Islands.Trans,Saoro nat,Hiat.Soc,, 13:85-102 1933 Mohr, N.,Forsg tuen Islandek Naturhiatoris. KjbenMvn 1786 Moign, A.,Contribution & l'etude littorale et sous-marine do la baie du Roi (Spitaberg, 79'N),Cab, 1965 0oanogr,, 17:543-63 Moore, P.G.,Ecological survey strategy,Mar.Pollui,Bull., 2:37-8 197 la The nematode fauna associated with holdfasts of kelp (Laminaria hy'perborea) in north- 1971b east Britain.J,Mar,Bi.ol,Assoc.U.K,, 51:589-604 FIRM s8Laninaria erborea Mowat, J.A,,Auxins and gibborellins in marine algae,Proc.Int.Seaweed ESy,np,, 4:352-9 1964 hrnda, I.,On Laminaria species from the Foslie Herbarium with special rogard to Laminaria intermedia 1965 Foslie.Nova Hodwigia, 9:285-90 Myklestad, S,,Experiments with seaweed as supplemental fertilizer,Proc,Int.Seaweed Symp,, 4:432-8 1964 ai, M., Marine algae of the Kurile Isl,1.J,Fac.Aric,Hokkaido (Imp,) Univ., 46:1-310 1940 Nansen, F.,Spitsbergen waters.Oceanographic observations during the cruise of the VESLMU! to 1915 Spitzbergen in 1912,Skr.Vidensk,Christiana (Mat.-Nat.), 1915(2):132 p. Newton, L.,A handbook of British seaweeds.London, British Museum, 478 p. 1931 Norton, T,A.,Underwater observations on the vertical distribution of al at St Mary's, Isle of 1968 Soilly.Br.Phycol,Bull., 3:585-8

Norton, T,A, and E.M, Burrows,Studies on marine algae of the British Isles,7,Saccorhizao 1969 schiden (Lightf,) Batt.Br.Phycol.Bull,, 4:19.-53 Ohini, H,, ist of common names of algae,Bull.Fac,Fish.Hokkaido Univ., 19:199-28 1968 Owen, E.C.,The carotene, carotenoid and chlorophyll conten$s of sorne Scottish seaweeds.J,Sci.Food 1954 Agrio., 5:449-53 Parke, M.,Studies on British Laminariaceas,1.Growth in Laminaria saocharina (L) Lamour.J,Ma. 1948 Biol.Assoo.U.K., 27(3):651-709 Parke, M. and P.S. Dixon,Check-list of British marine alre second revision,J.Mar,Biol.As000, 1968 , 48(3):783-832 Peat, S., W.J. Whelan and H.G, Lawley,The structure of laminaria.J,Chem.Soc,, 1958:724-37 1958 Percival, E. and LH, McDowell,Chemistry and enzymology of marine algal polysaccharides.London, 1967 Academic Press, 219 p. Percival, LG.V, and A.G. Rosa'The cellulose of marins algae.Nature, Lond., 162:895.-6 1949 Fucoidin,Part 1.The isolation and purification of' fucoidin from brown seaweeds. 1950 J.Chem.Soc., 1950(145):717-20 Prints, H.,Die Algenvege-tation des Trondhjemsfjordee.Skr,Nor.Videnek.Akad, (Ma-t.-Nat.), 5:1-273 1926 Reos, T,K,,Fruiting periode of' brown seaweeds.Proc.Swansea Sci,Field Nat,Soc,, 1:33-7 1928 Reid, K.C., Production of mannitol and laminaria from seaweed aqueous extracts,Proc,Int.Seaweed 1956 Symp., 2:105-6 ReidK.C. and P. Jackson,Non-thermal drying of brown marine algae.J,Sci,Food Agrio., 7(4)291300 1956 RUnnorstrand, S., Untersuchungen über Oxydase, Peroxydase und AscorbinsMure in einin Moerosal 1943 Lund, C.W. LindstrUm 9:10 FIF 58Laminaria rborea RoB9nvinge, L,K, and S. Lund,The marine algae of Denmark.Vol. 2.Phaeophyceae.3.Bioi.Skr,, 1947 4(5):1-99 Russell, 0,,Larninarioco ax tomentosoides on the Isle of Man,J,Mar.Biol.Aaaoc.U,K,,44(3):601-12 1964 The genua Ectocarpus in Britain, 1.The attached forms.J.Mar.Bioi.Assoc.U.K,, 1966 46(2):26794 Saelen, 0.H,,The hydrography of some fjords in northern Norway:Baisfjord, Ulefjord, Trphsund, 1950 Vorgsyfjord and Malangen.Promace Mui.Aarsh., 70(1947), 1:102 p. Sauvacau, C.,Recherches sur lee iriminaires dea o6tee de France,Mm,Acad,Sci,Inst.Fr., 56:233 p. 1 918 Schnepf, E.,Goigi.-apparat und Sekretbildung in den BrUsenzellen der Sohleimgunge von Laminaria hyper- 1963 borea.Naturwissenschaften, 50:674 Schreiber, E.,Untersuchungen Uber Parthenogenesis, Geechlechtabeetimmung und Baatiardierungsverm15gen 1930 bei Laminarien.Planta, 12:331-53 Schuitz-Schuitzenstein,Ueber Schichtenbilclung im Pflanzenreich mit Beziehung auf die natUrlicI 1853 Classification der Pflanzen,Flora, ,Yona, 36:49 Seoane Carnia, J.,Las la1ninarias da Eapaia y au distribuci6n.(me kelpa of Spain and their diatri- 1966 bution).Publ,Tda,Junta Eatu&Peaca, (5):425-36 Setchell, W.A. and N,L, Gardner,The marine algae of the Pacific coast of North America,Part 3. 1925 Melanophyceae,Univ.Caiif,Publ.,Bot,, 8:383-898 Seybold, A. and K, Egia,Quarrtitative Untersuchungen Uber Chlorophyll und Carotenoide dar Meez'eaaln. 1938 Jahrb.Wias.Bot,, 86:50-80 Sloane, J.F, et al.,The ecology of the Lough Ins rapids with special reference to water ourrente.5. 1957 The sedentary fauna of the laininarian algae in the Lough Ins area.J,Animn.Ecoi., 26(1): 197-211 Smfrholm, 0,,Maskinell h3sting av etortare.Tangmelpos'n, (8) 1964 South, G.R, and A, Cardinal,A checklist of marins algae of eastern Canada.Can,J,Bot.,, 48:2077-95 1970 Spence, M.,:Larninariacene of Orkney:Their ecology and economics.J,Bot., Lond., 56:281-5 1918 Strm, H.,Physiak og Oeconomiak Beakrivelse over Fogderiet Sndmere, beliggendei Bergerie Stift i 1762 Nor. Soros,Del i Sunde, I.,The iodine content of eggs from hens fed with seaweed meal,Proc.,Int.Seaweed Sno,, 1956 2: 109-13 Sundane, 0,,The algal vegetation of Osiofjozd.Skr.Nor.Vidensk.Akad., 1953(2);244 p. iq'

Interfertility between forms of Laminaria digitata,Ny-ti Mag,Bot.6:121-8

The ecology of Laminaria digilata in Norway in view of transplant expimenta.Ny-tt 1964 Nag.Bot,,11:83-107 FIRM/S87 Laminaria hyperborea :11 Svend.sen, P.,The algal vegetation of Spitsbergen.Skr.Nor,Polarinst.,, (116»49 p. 1959 Svendaen, P. and J.M. Kam,The taxonomic status, distribution and morpholo' of Laminaria cucullata 1971 sensu Jarde and Kiavestad.Sarsia, (46):1-22 Tokidá, J.,Marine algae epiphytic on Laxnizmrìaios plante.Bi.ill.Fac,Fish.Hokkaido Uair,, 11(3): 1960 73-105 U.K. Admiralty,Uydrograuhic Department,Admiralty tide tables, 1964.Vol. 1.European waters. 1963 London, U.S. Nav',Rydrographic Office.World atlas of sea surface temperatures.Second ocLition.Publ 1944 ydrogr,Off,, Wash., (225) Vahi, O.,Growth and deneity of Patina pellucjda (L.) (Gastropoda: Prosobranchia) on Laminaria 1971 hyperborea from western Norway. Ophelia,9 (in press) Vinogradov, A.?. (j. Efron and J,K. Setlov, Tranels) The elementary chemical composition of marine 1953 organisms,Mem.Sears Found.Mar.Res., (2:648 p. Walker, F.T,,Sublittoral seaweed survey of the Orkney Islands,J,Ecol,, 38:139-65 1950 Sublittoral seaweed survey:Dunbar to Fast Castle, EasScotland,J,Ecol., 40(1): 1952. 74...83 Distribution of Laxoinariaceae around Scotlanci,Nature, Lond,, 113:766-8.Issued also 1954 as J.Cona,irm.int,Explor,Mer, 20(1):16O-6 ,The Laminaria cycle. Rev0a]0, (3):179-81 1956a Periodicity of the Laminariaceae around Scotland.Nature, Lond,, 177(4522):1246 1956b ,An ecological study of the Leminariaceae cAilsa Craig, Holy Island, Inchmaa'nock, May 1957 Island and Seaforth Island.Trans.Proc.BotSoc.Edinb,, 37(3):182-99 Walker, F.T, and W.D. Richardson,An ecological investigation of Laminaria cloustoni Edm, (L. hyper- 1955 borea Foel) around Scotland,J.Eool,, 43:26-38 Perennial changes of Laminaria cloustoni Edn, on the coasts of Scotland.J,Cons.Psrm, 1957 Inb,Explor.Mer, 22:298-308 Whittick, A.,The kelp forest ecosystem at Petticoe Wick Bay mt, 55°55'N long. 2°O9'W an ocological 1969 study.Theeis, Durham University, 139 p, Wilce, R.T.,Studies in the niis Laminaria.1.Laminaria cuneifolia J.C. Agardh:a review, 1959 Not., 112(2):158-74 Studies in the genua Laminaria.3.A revision of the North Atlantic species of the 1965 simplices section of Laminaria,Bot.Gothoburg., 3:247-256 Williams, J.L.,The gametophytes and fertilization in Laminaria and Chorda. AxLfl.Bot, 35:603-7 1921 9,12 V.EPJA/887Laminaria hyperborea

ADDENDUM

Bellamy, D.J. and A. Whittick, Problems in the assessment of the effects of pollution on inshore 1968 marine ecosystems dominated by attached macrophytes. In The biological effects of oil pollution on littoral communities, edited by J.D. Carthy and D.R. Arthur, London, Field Studies Council, pp. 49-54

Ernst, J. Quantitative data of the vertical distribution of Laminaria along the coast of Rretagne. 1966 C.R.Hebd,Séances Acad.Sci.Paris, 262 (26):2715_7

Fleming, M., Sir Edmund Hirst and D.J. Manners, The constitution of laminarin 6. The fine structure 1966 of soluble laminarin. Proc.Int.Seaweed Symp.5, 255-60 Mainel, G., Phéophycées de France. Paris, 1931-1939. 431 p. 1938

Hoek, C. van den and M. Donze, Algal phytogeographyof the European Atlantic coasts. Blumea, 15:63- 1967 89

Lamouroux, J.V,, Essai sur les genres de la famille des Thalassiophytes non articulées.Ann.Mus.Hist. 1813 4at., 20(1)21_47

Munda, I., Differences in the algal vegetation of two Icelandic fjords, Dyrafjbrur (West Iceland) and 1969 Reyùarfjbr&*r (East Iceland). Proc.Int.Seaweed Symp. 6: 255-61 SYNOPSIS OF FISHERIES BIOLOGICAL DATA

Thisis one ofaseries of documents issued by FAO, CSIRO and USFWS concerning species and stocks of aquatic organisms of present orpotential economic interest.The primary purpose of this seriesis to make existing information readily available to fishery scientists according to a standard pattern, and by so doing also to draw attention to gaps in knowledge.Itis hoped that synopses in this series will be useful to other scientists initiating investigations of the species concerned or of related ones, as a means of exchange of knowledge among those already working on the species, and as the basis for comparative study of fisheries resources.They will be brought up to date from time to time as further information becomes available either as revisions of the entire document or their specific chapters.

The relevant series of documents are: FAO Fisheries Synopsis No. FR/S replacing, as from 1.1.63, FAO Fisheries Biology Synopsis No. FB/S CSIRO Fisheries Synopsis No. D FO/S and USFWS/FAO Fisheries Synopsis No. N M FS/S

Synopses in these series are compiled according to a standard outline described in Fib/Si Rev. 1(1965). FAO, CSIRO and USFWS are working to secure the cooperation of other organizations and of individual scientists in drafting synopses on species about which they have knowledge, and welcome offers of help inthis task. Additions and corrections to synopses already issued will also be most welcome. Comments including suggestions for the expansion of the outline and requests for information should be addressed to the coordinators and editors of the issuing organizations. FAO: Fishery Resources Division Marine Biology and Environment Branch Food and Agriculture Organization of the United Nations Via delle Terme di Caracalla 00100 Rome, Italy USFWS: CSIRO: Department of the Interior Scientific Editor Bureau of Commercial Fisheries CSIRO Division of Fisheries and Oceanography Office of Scientific Publications Box 21 Building 67, U.S. Naval Air Station Cronulla, N.S.W. Seattle, Washington 98115, U.S.A. 2230 Australia

Consolidatedlistsof species or groups covered by synopses issued to date or in preparation will be issued from time to time.Requests for copies of synopses should be addressed to the issuing organization.

The following synopses in this series have been issued since January 1970:

DFO/S4 Synopsis of biological data on the rainbow prawn, Parapenaeopsis sculpt//is (Heller, 1862) 1970 DFO/S5 Synopsis ofbiologicaldata on the schoolprawn, Metapenaeus mac/eayi (Haswell, 1879) 1970 BCF/54 Synopsis of biological data on chum salmon Oncorhynchus keta July 1970 F1RI/S80 Synopsis of biological data on the eel Anguilla anguilla (Linnaeus, 1758) October 1970 Rev. i * FIRM/S91 Synopsis of biological data on Cran gon crangon October 1970 * FIRM/S92 Synopsis of biological data on Panda/us montagui October 1970 * FIRM/S93 Synopsis of biological data on the Jumbo tiger prawn Penaeus monodon Fabricius, 1798 October 1970 * FIRM/S94 Synopsis of biological data on the Indian prawn Penaeus indicus H. Mime Edwards, 1837 October 1970 * FIRM/S95 Synopsis of biological data on the prawn Panda/us platyceros Brandt, 1851 October 1970 * FIRM/S96 Synopsis of biological data on the penaeid prawn Solenocera indica Nataraj, 1945 October 1970 * FIRM/S97 Synopsis of biological data on the penaeid prawn Metapenaeus dobsoni (Miers, 1878) October 1970 * FIRM/S98 Synopsis of biological data on the penaeid prawn Metapenacus affinis (H. Mime Edwards, 1837) October 1970 * FIRM/S99 Synopsis of biological data on the ocean shrimp Panda/us Jordan! Rathbun, 1902 October 1970 * FIRM/Si 00Sinopsis sobre labiología del camarón blanco Penaeus schmitt! Burkenroad, 1936 October 1970 * FIRM/Si 01 Synopsis of biological data on the white shrimp Penaeus setiferus(Linnaeus, i767) October 1970 * FIRM/Si 02Synopsis of biological data on the brown shrimp Penaeus aztecus aztecus (Ives, i89i) October 1970 * FIRM/S103Synopsis of biological data on the pink shrimp Penaeus duorarum duorarum Burkenroad, 1939 October 1970 * FIRM/Si 04 Synopsis ofbiological data on the penaeid prawn Metapenaeus monoceros (Fabricius, 1798) October 1970 * FIRM/Si 05Synopsis of biological data on the penaeid prawn Metapenaeus bray/corn/s (H. Milne Edwards, i837) October 1970 * FIRM/Si06Synopsis of biological data on thepenaeid prawn Parapenaeopsis sty//bra (H. Milne Edwards, 1837) October 1 970 * FIRM/Si 07Sinopsis sobre la biología del camarón nailon Heterocarpus reedi Bahamonde, 1955 October 1970

DFO/S6 Synopsis of biological data ori the greentail prawn Metapenaeus bennettae Racek and Dall, 1965 1970 DFO/S7 Synopsis of biological data on the eastern king prawn Penaeus plebejus Hess, 1865 1970 DFO/S8 Synopsis of biological data on the banana prawn Penaeus mergu!ens/s de Man, 1888 1970 FIRM/S82 Synopsis of biological data on North Atlantic sandeels of the genus Ammodytes (A. tobianus, A. dubius, A. americanus and A. marinus) November 1970 FIRM/S83 Synopsis of biological data on Saccorhiza polyschides October 1970 NMFS/S79 Synopsis of biological data on Pacific Ocean perch, Sebastodes alutus December 1970 FIRM/S38 Synopsis of biological data on knobbed wrack Ascophyllum nodusum (Linnaeus) Rev, i Le Jolis December 1970 FIR M/S84 Synopsis of biological data on haddock Melanogrammus aeg/ef/nus (Linnaeus, 1758) December 1971 FRm/587 Synopsis of biological dota on Laminaria hyperborea December 1971

* Published in the Proceedings of the World Scientific Conference on the Biology and Culture of Shrimps and Prawns, Mexico, 12 to 21 June 1967.

MI/C3845/2.72/E/1 /900