Organic Aggregates Formed by Benthopleustophyte Brown

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Original Citation: Organic aggregates formed by benthopleustophyte brown alga Acinetospora crinita (Acinetosporaceae, Ectocarpales) / Giani, Michele; Sartoni, Gianfranco; Nuccio, Caterina; Berto, Daniela; Ferrari, Carla Rita; Najdek, Marjana; Sist, Paola; Urbani, Ranieri. - In: JOURNAL OF PHYCOLOGY. - ISSN 1529-8817. - ELETTRONICO. - 52(2016), pp. 550-563. [10.1111/jpy.12413]

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08 October 2021 This article isprotected rights by All copyright. reserved. 10.1111/jpy.12413 differencesofversion Record. and c theVersion Please between this throughcopyediting,typesetting, the proofreading pagination and process, whichmay lead to This article undergone has for and buthas accepted been been not review publication fullpeer Articletype : Regular Article Accepted Date 07: Revised Date : 24 Received Date: 03

AcceptedAgenzia Regionale per Protezionela dell’Ambiente dell’Emilia Romagna, Article N Organic aggregates formed by benthopleustophyte brown alga brown by formed benthopleustophyte aggregates Organic ational Instituteational of Oceanography andExperimental Geophysics (OGS), 34151 Trieste, Italy Istituto Superiore per Protezionela e la Ricer A Institut “Ruđer Bošković”, Center for Marine Research, cinetospora crinita cinetospora - - Department Life of Sciences, University of Trieste, 34127 ItalyTrieste, - Feb Dec Mar - Department Biology, of University Firenze, of 50121 Firenze, Italy 15 - - 2016 - - 2015 2016 248

G ianfranco Paola Sist

C ( Marjana Najdek A M arla

D Sartoni, C cinetosporaceae, cinetosporaceae, aniela ichele , Ranieri Urbani

R ita

Berto Giani Ferrari ca Ambientale,ca aterina

Nuccio 2

52210 E 30015

ite this article as ite this doi: ctocarpales) Rovinj, Croatia 47042 Chioggia,Italy Cesenatico, Italy

This articleThis protectedis by copyright. All rights reserved. Organic(Dissolved Organic Nitrogen), DOP Phosphorous) Esters),(Polyunsaturated PUFA Fatty Acids), TCHO (Total carbohydrates), DCHO (Dissolved carbohydrates), FAMEs(FattyAcids Methyl Abbreviat polysaccharides Key heterotrophicgrowth. to the fattyacid likely quite low,most was pool for tothe knownpoortheir due conditions crinita The fatty for patterns acid theaggregates found were tothoseobserved similar inthe isolated exopolymer resembledcompositions theaverage ofthecomposition Tyrrhenian high molecular weight mucilages alga produced by brown ofinside/outside theaggregates. The and Ndisplay their originand marine evidence could ofthethatoccur provide processes the development phytoplanktonabundant of an community. stableisotopesof C Analysesofthe which regeneration of nutrientsandorg recycling dissolved of the of bottoms Mediterranean Sea formed bybrown the filamentous alga This work presentselemental, polysaccharide, the and benthic fattyof compositions acid aggregate Abstract Responsibility:Editorial Posewitz M. Editor) (Associate

Accepted index Article

algae withvariablecontributions from diatomspecies. embedded b The

ions: ions: words: s , a nd the observed differences couldnd theobserved beascribed differences todifferent degradation stages. CTD probe (Organic (Conductivity,CTD OM probe), Temperature, Depth Matter), .

Acinetospora crinit

The aggregatescancharacterizedThe as be mineralized A. crinita A.

monosaccharide monosaccharide a Acinetospora crinita, crinita, Acinetospora ,

aggregates, brownalgae, algae,filamentous

were quite similar. Inwere quite similar. particular,the sample Adriatic DOC (Dissolved OrganicDOC (Dissolved Carbon),DON

compositions .

which are spread ontherocky widely anicactively matterand favor occur

of Adriatic and Tyrrhenian acterial contribution lipids, centers

s This articleThis protectedis by copyright. All rights reserved. the main exopolysaccharidic componentsfraction. ofthe 26.6% to55.9%of differentfilamentousal. that c algae (Sartoni2008)showed et the benthic communities. causingsettle ontotheseafloor conditionsof thatca and theformationanoxic detachaggregates the macroalgae thefrom they are usuallyattached, towhich allowing to them bleaching 1995) byofcausing gorgonians suffocation ( (Innamoratial. et 2001,Misic al.2011). et TyrrhenianLigurianandareas where mucilaginous thebenthic aggregateswere observed (Erikssonanoxia et al.1998and ofeutrophication no referencesevidence exists inthe therein), an increased occurrence ( algae, species this is als et al.2000 Tyrrhenian ( Sea frequentlyobservedmu reported inthe benthic taxa 1992 belonging Ectocarpales, tothe order inthe Mediterranean iswidespread ( Sea 2 1 Sartoni and1992 Sonni

Received December xx 2015. Accepted xx Author for correspondence: Previous studiesonthe chemical ofbenthic composition aggregates andbiochemical formedby Mucilaginousaggregatescan (Calvo benthic theetal. adverselybenthic impact communities Although inlocally studies Baltic eutrophicated coastalthe areas Sea allover region shown have The free ,

Accepted Article Welker and 1994 Bressan

of of ) and Ligurian) andSea 2007). (Schiapparelli Together with other filamentous etal. brown - living formliving of corallin Sartoni and 1992 Sonni Sartoni the organic carbon, respectively. mannose Galactose, xylose or and were fucose ales or scleractinians (Schiaparelli currents bottom can etal. 2007).Moreover, o largely throughout intemperateglobe distributed o coastal the environments , of fast

Pedersen 200 Kristiansen and Acinetospora crinita Acinetospora email - growing filamentous algaegrowingfilamentous and loose ,

Antolić :

[email protected] ,

et al.2010, Innamorati 1995,

Mistri andCeccherelliMistri 1996

(Carmichael Harvey)a ex Sauvageau, cilaginous inthe aggregates examined , phone +390405588762

, Tsiamis etTsiamis al.2013 1 ).

Mistri andCeccherelliMistri 1996 arbohydrates and proteins accountfor arbohydrates - lying algallying matsthat cause , ) and is one is ) and ofthe most Giuliani etal. 2005)or

Sartoni and Sonni n adversely impact

brown alga , Hoffmann This articleThis protectedis by copyright. All rights reserved. sedimentation chambers. weremagnification at on subsamples using performed 400X Zeiss a mL) withneutralized formalinfixed concentration werea to of final 4%. observations Microscopic pe performed macroalgal todescribeandMacroalgal the microalgalcompositions. observations were Microscopy analyses Temperature, Bird Depth, Electronics SEA ultrapure Milli immediately at frozen aggregatesfor were different dividedeleme analyses.For 2004byand 26June divers scuba L). polymethacrilate using Land 2 (0.06 The syringes sampled Sampling andphysical parameter measurements peninsula (Croatia)and inthe southern sub sub two areasSea, thei.e.,inNorthern Adriatic of Site descriptions MATERIALS METHODS AND allowing thus bottom, a better understanding theirmarine origin environment. andfate of inthe aggregatesaggregates fromal. (Giani 2012andreference pelagic that et sediment onthe therein) ofbenthiccomposition aggregates formedby different distingu to algaeisfundamental benthic aggregates inother Mediterranean found areas. chemical Knowledge ofthe andbiochemical f of the organic matrix rformedat For 250Xmagnification. of themicroalgalcomponents, identification samples(250 Accepted Salinity, Article The tocharacterize aim aggregates ofthisstudy thebenthic was andwhether nature toassess the

temperature, and depth profiles and temperature, werea determinedusing probe CTD - Q water, and finallyQ water, freeze . Sampl . Light of microscopeexamination thecollected aggregates benthic was ormed by algafilamentous brown the – 20°C, successively20°C, de ing aggregateswas ofbenthic 2004in 2 and carried 26June outbetween

- - dried. - basin near basin the Tremiti

SBE 25). SBE - frozen against 3500MWCO) and dialyzed(Cellusep, .

The benthicwere aggregatescollected between 2

basin along the western coast of the Istrabasin alongthewesterncoast ofthe ntal analyses, the subsamples werental analyses, the A. crinita A. Islands (Fig IM35 invertoscope in

is similar to that ofthe .

, (Conductivity, a and b) ishing these

This articleThis protectedis by copyright. All rights reserved. potassium nitrogenand total dissolved colorimetric methodsLuebbe withaBran Autoanalyzer Total Mod.AAIII dissolved instrument. treatment. reproducibilitywas <0.1 Belemnite) analytical The standards. measurement was of the precision 0.2 The( results asmil were deviation expressed per Isotope Mass Spectrometer Ratio withaCHNS coupled 1 spectroscopy (ICP Canada)(CNRC, was toassessof the accuracytheanalytical used data. was analyzer Fisons as paper reporteda 1108EA, The previous et (Sartoni reproducibility al.2008). in by onignition loss of 100 weight drying after 48hatThe loss for 60°C. matter organic gravimetrically (OM)was determined Elemental analyse andisotopic macroalgal andtheircomposition relative abundance by viathe methodsuggested (1968) Jones 976). The The determination ammonia, nitrites, phosphates of nitrates, and silicates were out via carried Stable carbon ( contentdeterminedPhosphorus was emission byplasmaatomic inductivelycoupled Total carbon Light ofthe microscopeexaminations collectedaggregates todescribe weretheir used 

Accepted Article3% forcarbon,

accordingKoroleff(1983 a, tob).

accuracyreproducibility ±4.8%and andwere 98.8 (C  - tot AES) at 177 nm on 1 mol at on1mol 177nm AES) 13 ), nitrogen (N ), nitrogen C) and nitrogen andC) (  5% forand5% nitrogen ‰ - 200 mgaggregates of after combustion 450°C. for 4h at . The phosphorus were by determinedafterwet persulfate oxidation s .  Water content determinedgravimetric was by measurement of 13 tot C valueC was determined afterthe re ) and sulfur (S) were) and(S) elementalCHNSO sulfur determineda using  15 N) isotopeswereanalyzeda using Finningan Plus Delta 

1% for1% sulfur, and the . L - 1 ‰

HCl extractsHCl ofthecombusted ) from the international VDB) from theinternational Dee (Vienna Pee

- O AnalyzerFisons (Italy) model  5%, respectively. reference mat moval of carbonates via HCl ofcarbonatesmoval viaHCl

‰,

and the sample (Aspila erial BCSS erial BCSS

EA1108.

This articleThis protectedis by copyright. All rights reserved. chromatography a with a(GLC) 6890NGC system5973NetworkMass in equipped Sele Morrison (1964). Fatty andSmith (FAMEs) wereesters methyl acid byAgilent analyzed gas me HCl), Fatty acid analysis was 1.5%. filmthickness)20 independent30 m×0.25mm×0.2μm and detector. of an RSD FID analyses The chromatograph Autosystem(Perkin Elmer Inositol16 h. aninternalas T was standard. used (2004), after polysaccharide hydrolysis in method of Blakeneyal. alditol acetates, (1983) et foras of described preparation ofexopolysaccharidescomposition via gasusing analyzed chromatography was thederivatization purification metho andpurificationIsolation ofmucilage polysaccharides calculatedglucose standard. basedonthe forhydrolysis at100°C 16h.The with0.1NHCl carbon ofthecarbohydratescontent was 2 analysis. samples were at stored pore WhatmanGF/Ffor filters) size, carbohydratesextracellular analysisdissolved of All (DCHO). analysisgently total carbohydrates andremainder of was (TCHO), filtered(0.7 the inner insidethe water aggregate, into was which divided Carbohydrate analysis. - benzothiazolinone hydrazonebenzothiazolinone hydrochlorideassayand (MBTH) Sieburth (Johnson Accepted ArticleTotal anddissolved carbohydrates were thylatedBF3(14% and extracted inmethanol), indichloromethane as reported

ds were described et inthe(Sartoni 2008paper al.2008). . Mucilage sampleswere acidified(6 M (1.2 saponified MNaOHinmethanol), -

Carbohydrates bycollecting sampled 200mL were ofbulk and seawater 20°C after0.02% the NaN20°C addition of

2 N trifluoroacetic acid (TFA) and heating2 Ntrifluoroaceticacidand at for (TFA) 100°C XL) XL) determined spectrophotometricallydetermined the3 using equipped with a capillary column SP2330 (Supelco, SP2330 with aequipped capillarycolumn he derivativesanalyzedhe wereinacapillary gas

. The exopolysaccharide extraction and two aliquots. Onetwo aliquots. aliquot was usedfor 3

as an antibiotic preservativeantibiotic as an until

The monosaccharide by et Magaletti al. - mm nominal mm 1977) after by ctive - methyl - l iquid iquid - This articleThis protectedis by copyright. All rights reserved. werethermohaline whereas properties inthefollowing quite homogeneous, a period, s increasing AtthebeginningfromIstra, trend February/March toJune. thewater in ofMay Thermohalin RESULTS growth, ratio this isestimated et as(White al.1980). >2.3 ratios wereC15:br/C15:0 used asgrowth bacterial indicatorsaggregates; for the for within optimal tototal C18:1n7) of + theprincipalmarkers +C17iso C17anteiso + C15anteiso + bacterial+ (C15:0C17:0 +C15iso mucilage. withrespect diatombiomass the relative of to proportion activity). ratios (Leveille C14:0+C16:1+C16P/C16:0 Diatom were et applied al.1997) todetermine determinegrowth stage the and were andC16P/C18P from ratios. C16:1/16:0 theThe fatty contributions ratios acid use recognizedaggregates, phytoplankton inpelagicconditions of growth the andbacteria were inferred thetotalfattycontribution to content acid all mucilaginous in aggregates. mucilaginous toassess thesolely fatty matrix acidalgae composition totheits and due hence pureindividual ofFAME standards also were used. C18 data for GCstan ultra 0.25 0.3mm Detector, crosscapillaryx (25μm; mx column The relativeof bacterial importance activity aggregates estimated from was inthe contributions To reveal microalga possible In thenativeaddition to benthic ag byThe and FAMEs a identified familyofequivalentchain were massspectralplot data length - - C20, polyunsaturatedC20, fattyacids (PUFA1liver oiland standards variouscod and PUFA3), Accepted Article high asthe carriergas. purity helium

e conditions dards for the specific GC column. The bacterial FAME standard mix, FAMEs mix FAMEs thespecific for The mix dards mix, GCbacterial standard column. FAME fatty themucilage in acids etet(Mayzaud al.1989,Najdek al.2002). The . Theand surface near l condition ofthe aggregated(as phytoplanktonameasure oftheir - bacteria interaction within thebenthicwithin bacteria mucilage interaction aggregates, as gregates,

- A. crinita bottom bottom

temperatures (

other phytoplankton classesother inthe phytoplankton algae from were the also isolated

- linked 5% and phenylmethylsiloxane) Table 1) Table

showed a similar trong d to This articleThis protectedis by copyright. All rights reserved. material the benthopleustophyticexclusively appears form tobe constructed from of AcceptedTyrrheniancounterpartsbased is percentage onthelower of aFrom ofview,peculiar point aspect this ofth (Aktan and Topaloğlu 2011) is actuallyTyrrhenian tothe for limited Sea species were recently introduced Tyrrhenianaggregates. lastaspect surprising not This is considering these twoallochthonous that Chrysophaeum aggregates, chrysophytes both chrysophyte theTyrrhenian usuallypresent in aggregates. Furthermore,inthe Adriatic benthic species with isoften mixed their development, alsoBressan Welker by and asnoted and(1994), th both cases, thefree that theyare s mucilaginousaggregatesas benthicsampledinthetwosites ofthe ArticleMacroalgae identification the seasonal spring warming. of riversgradient Athermal area. inthestudy decreasing from theto surface Islands,site. AttheTremiti the salinity was (38.48 high located between decreasedsurfaceIstrian thesalinity of along waters withahaloclinecoast the 36.1, downto the temperature 16.0 ranged from betweenand thethermocline and 22°C 23°C, isl IstralatitudeIn ofthe Istriantemperature peninsula. coastal inthe thesurface ranges waters June, stratificationIsland, was AtTremitiwarming than at established. the was relevant the higher more imilar to those present areas tothose various oftheimilar in Tyrrhenian(SartoniIn Seaet al.2008).

taylorii – 4 ma - living form of living form

nd nd I.Lewis F. . – The ofthe observations macroalgalthat produce community the Nematochrysopsis marina 8 matSecca Croce Punta and Porer andBanjole slightlyatdeeper the

for

Chrysonephos lewisii C. taylorii C. into theMediterraneaninto and et their al.1995), distribution (Sartoni °C

A. crinita et

H. F. BryanF.were H. absent, although found they inthe were

16.5°C. At 16.5°C. the enda of water freshened May, input .

C. lewisii C. (Carmichael) afundamental Sauvageau plays in role e aggregates Adriatic compared ocated betweenocated

( W. R.Tylor)W. and R.Taylor W.

and tothe Tyrrhenian and Aegean Seas (Feldmann) filamentous Billard,a - 38.77), indicating no significant38.77), indicating influence N . marina. - 10 m and and 10 m Adriatic SeaAdriatic (Fig is benthopleustophytic

Most ofthe examined Most - 20 m; below line, 20 m; this the bottom wasthe bottom due to

with their .

A. crinita, A. 2) have shown 2) shown have

both This articleThis protectedis by copyright. All rights reserved. et al.2001 previouslyevidenced previousof from observations theTyrrhenian benthic theAdriatic aggregwithin benthic Microalgal community (Athanasiadis 1997 Mediterranean and negativel materialthis an is allochthonous filamentous Rhodophyta inthe entire that widespread is macroalgae, substrate of for benthic mucil thedevelopment and vertical are largelyovergrown surfaces by turf Islandsgorgoniansare and thecoralligenous in communities, lacking assemblages onhorizontal ephemeralIn bushy thecircalittoral ofthe macroalgae. examined most stations zone, and Islandsappears rathe asFrom ofview,noted point this circalittoral of gorgonians thecommunities represent zone, for asuitable their support development. perennialgenera algae belonging brown tothe accumulation infralittoral inthe lower particularly zone, such anchoras inthe presenceof species of These quite understandable differencesiftheeven benthic are not aggregates ofthe free aggregates.compared site withthe Rovinj different becausea laterappear month and they evolution temporal showremarkably early aging during the middle until from of theend thesurveys performed April butt July, of betweenIsland Tremiti thetwosites.aggregates The benthic only not show in the aggregates col

A. crinita AcceptedSargassum Article ,

Sartoni et al. 2008). Abundances varied from2008). Abundances 10 Sartoni et varied al.

areusually inoccurrence. these unpredictable Nevertheless, showa materials massive Womersleyella setaceaWomersleyella

spp. are present,the sporadicgrow andtherefore, not aggregateson primarily , Piazzi etal. 2007 Piazzi Islands, and inthe Tremiti inRovinj lected exists ifa even difference striking r unstructured; the infralittoral photophilic communitiesr unstructured; the infralittoralphotophilic with . Both at Rovinj and the Tremiti Islands,Both and theTremiti atRovinj living themicroalgal community yaffects and number diversity thespecies ofinvaded

by Cormaci(2001), thebenthic etal. vegetation ates composed was as ofmicrophytobenthic diatoms, largely

(Hollenberg) R.E. NorrisR.E. (Hollenberg) particularly appears tobe abundant; , Nikolić et al.2010

aginous aggregates. Among the turf aggregates.aginous Among Cystos - formingwhichan represent unsuitable species,

eira

). and

to 10 Sargassum 7

cells . , whereas inthe L

- aggregates (Innamorati 1 poor development , and a shifti time hey a also exhibit

of theTremiti communities communities - Cystoseira forming

in the Tremiti - living fo living

spp. n rm This articleThis protectedis by copyright. All rights reserved. OM/organic showed carbon ratio an of2.26 average value 91.6 Elemental theaggregates composition of closterium comparable Rovinj the tothatof but was usedonlycompareAt to selectedsamples. Theof scarce thesporadicaggregates atthe Tremitinotdepict sitesdid trend developmentasimilar Grammatophora, Synedra accompanied diversified by aand diatom community( rich together with nano Cylindrotheca closterium showed notably lowabundances onthe mucilage spp. and microphytobenthos substratescollected atnotaffected the sameon by depth mucilages. period, performed a was comparison thecommunity between and aggregates ofthe a thecate benthic In worth samples, itis noting these thepresence of ( Pleurosigma, Amphora, Grammatophora, Cocconeis, Ist together ( withdiatoms a many higher presencepennate of Reimann (Fig.3).From the unipunctata Striatella the dominantdiatomgenus, rangingoverfrom 80% total abundance, by 45%toofthe followed of composition taxonomic theaggregates Rovinj wasInasMay, revealed well. Dictyocha fibula In June, Accepted% Article ;

Table 2 Striatella unipunctataStriatella

was detected not Cylindrotheca closterium ) with an organic) with ancontent matter thatrangedto63%.The (OM) from 41%

dinoflagellate DSP

Ehr -

and micro and enberg

(Lyngb

reached percentage a higher (31%) ); it become); it component of the theaggregates nearly exclusive inJuly.

beginning to the end of May,beginningthese decreasedend thedominancetaxa of of tothe

, Rhabdosphaera claviger ye - , which dominated in the aggregates41%,, which respectively), dominatedand (47%

unidentified diatoms. pennate ) Agardh and aggregates3), (Fig. -

producing species. For the aggregates collectedproducing For theaggregates during species. this appeared abundant most and species tobethe was always . The Cylindrotheca closter Cylindrotheca A crinita A - free respectively),and (4% 3%, substrates whereas

Prorocentrum lima the end was composition of May, themicroalgal

and in June, thedominanceand inJune, of

MurrayBlackman,& thecate dinoflagellates).

aggregates were hy highly hmia  Synedra, Navicula, Nitzschia, Licmophora, 0.68. The embedded carbonates inthe

with respecttheaggregates to (8%), ) and ofother specimens algalclasses ium

(

Ehrenberg

(Ehr Diploneis, Diploneis, enberg Licmophora ) Dodge,) which is drated (88.5 ) Lewin) Cylindrotheca Cylindrotheca Licmop et

spp. was spp. was

- hora

This articleThis protectedis by copyright. All rights reserved. reefFlorida in Faganeli et al and plankton( previous works for (average: to generallyas ofthe indicator terrestrial used (δ aggregates sampledinTyrrhenian seaby Sartoni degradation (fresher)ratios stages ifthese for are compared withthose reported thebenthic of (Del P Neg phosphatase intherespect aggregates with thesurroundingbe to thecause waterscould ofdepletion predominance P of of rigida phytoplankton (Redfield Redfield ratio comparableinmacroalgae 1963) andfound tothose 2008). the values aggregates inthe reportedAdriatic for andsea(Sartoni sampled et benthic Tyrrhenian al. northern Adriatic Sea e (Giani ranged values tothosereported 1.05%similar mucilagefor to2.23%, from sampledinthe pelagic ( aggregatesnon constituted a Table 2

- the Tyrrhenian aggregates benthic compared wit The The higher C The C 15‰) origin oforganicbenthic matter evidenced 1992), aggregates (Druffel& in Williams

Accepted Article

and A. crinita ), likely d org - 19.7 Fucus virsoides /N and C . ro et al. 2005). ro et al.2005).

, – (1988) for the macroalgaefor(1988)  less negative values ( less negative 20.7‰, Giani etal. 2009)samples. 20.7‰, org 1.4

sampled at Tremiti showedsampled at avalue mean of ue to the entrapment of organisms with calcareous with entrapmentorganisms Nitrogen uecontent skeletons. tothe of /S ratio values/S aggregates observed inthe benthic a couldresult beofdifferent - pelagic aggregates ( pelagic ‰; depleted organic compounds as polysaccharides. organicdepleted as compounds org /P /P

Fig molar ratiosaggregates ofthe

. (Faganeli et al. 1988). The(Faganelial. relatively C 1988). et higher

4a - negligiblefraction of theinorganic

t al. 2005). Sulfur content wast al.2005). Sulfur content relatively low( ) ,

showed a showed  13 - C: −16.6±3.1 C: 19.7 Ulva rigida

prevalence observed ofmarine origins, asalso in  13 2.2‰, Giani etal 2.2‰, C fromC

and coworkers (2008).

h those ofother typesaggregates of suggest a

( ‰ Fig. 4b - (Table 2) were higher , Lamb, etal. 2012) 30‰ to ), whereas algae inturf ona coastal δ 13 .

– 2006; C similarC tothat reported by constituents oftheaggregates 25‰) or marine25‰) (δ or Thealkaline activity high of - 19‰, Faganeli et19‰, al.2009) The δ The org have reported been /P molar ratios/P ( Table 2)

than the than 13 C ratio,C which is 13

with respect to C fromC Fig. 4a Ulva - . 23‰

) This articleThis protectedis by copyright. All rights reserved. in the wateraggregates. outsidethe between was water, and DCHO noted inthe interstitial DOC whereas noco higher. times concentrationwas value six that enrichmentnutrients. Moreover,DOC was four upto the aggregates (Fig. theouter to bulkwater 5 withrespect High of concentrations ( organicand nitrogen phosphorus respect the to silicates, nitrates and o waters Interstitial (Lassauqueetand al.(2010) authors therein). the seagrasses thearea sampled subjectrespectsites inputwith in topristine via anthropic anthrop 2012). relatively lowvalues (+2.7±0.9) ( 15 aggregates. signatureisotopic sampledIstra atin different the might sites periods suggest sampling achange elemental and of Vizzini et al.2003 Vizzini N signaturelowermacroalgae range fallsi.e., 0.5 inthereported by for other studies, Accepted ArticleThe analysis results ofnitrogen isotopic of The The negativerelationship b

 ogenic impact in the samplingogenic fact, impact sites,and of in in enrichment 15 N isotopic ratioN isotopic ofthe benthicranged aggregates from 5.29 2.68‰to

surrounding water (

with aging and degradation progressive bacterial , f the benthic aggregates Cole et al.2004) or4.5 ammonia, were interstitial waterammonia, intheaggregates’ concentrated highly with DOC and DCHO ( DOC etween were reportedturf for algae F F ig. 5 ig.

ig. 5c).  13 , a and b). Similar behavioraSimilar and b).was observ C foundC inthe 47.8±10.3 µM) were µM) 47.8±10.3

- . 10 ‰ All of the dissolve All of  A linear (n correlation = 5,R

15 N in thebenthic suggest aggregatescould N in

(Moncreiff and 2001); however, Sullivan

- fold higher, whereas the DCHOfold higher, reacheda whereas A. crinita , c and d),c as

on a reef coastal inFlorida(Lambal.on et d n found

and theC utrients, i.e.,phosphates, (Coffin 1989) ofthes(Coffin

in the interstitial water interstitial in the within

15 observed for theother N values was observedin org 2 rrelation was observed =0.952, p<0.01) /N ratios /N ed for theed dissolved for ‰ (Table 2). - (Fig. 4c),, 13.8‰ e a l

This ow

This articleThis protectedis by copyright. All rights reserved. fattyacid ratios were proportions and found in tothose similar AcceptedsamplesIstraIslands collectedalong The(Punta the andTremiti ranges coast Ferraio). for themajor Khotimchenko 1998). aregood data algae with agreement reported in other for (Fleurence from regions brown et al.1994, C18:4(n C18:1(n mucilage C16:0, C14:0, sampleswere Fatty acids xylose fucose and highergl of percentages also for shown Marked comparison. to becan notedwith respect differences the composition ofthe exopolysaccharideaggregates ofthe produced by includedcomposition glucose, rhamnose and(approx mannose (approximately followed 30%) by(approximately xyloseThe and remaining fucose 20%). showed similarmonosaccharide with patterns, Articleresults are shown averaging all Islands,Istra samplesfrom over and separately the coastfrom theand Tremiti the The mean the monosaccharide of compositions exopolysaccharidewere fraction by obtained Rib, ribose; Ara, arabinose; Xyl, xyl fucose; Fuc, rhamnose; Rha, mannose; Man, glucose; Glc, galactose; Gal, Abbreviations: percentages are (w/w) reported and arabinose) The were as monosaccharideexpressed compositions determined. relative formed by Carbohydrate characterization - 3)), and (i.e., PUFA C20:4(n C20

A. crinita . The observed acids major inthe fatty (FA) in (Fig. 6).

Fig neutral aldoses (galactose, glucose,neutral aldoses mannose,rhamnose, (galactose, xylose,ribose fucose,

The charThe ure ucose (32%) and galactose (32%)and (41.5%)ucose ofmannose, and percentages lower

6 .

The exopolysaccharidesThe produced by . acteristic features of acteristic features of ose. in The purified polysaccharides the mucilagefrom extracted samples Table 3

6) and C20:5(n and 6)

- galactose monosaccharide as themain 9), C18PUFA (i A. crinita

Acinetospora crinita - 3)), as reported in

wererecognized inbenthicmucilage imately 7 A. crinita .e., C18:2(n

A. crinita A. - . One dif N. marina N. 11%). In figure,11%). thesame -

6), C18:3(n isolated from benthic A. crinita Table 4. in these twosites ference( inalga

(Istra site)is

These resultsThese samples with - 3), A. A.

This articleThis protectedis by copyright. All rights reserved. Tanaka 1999). et al. 1988,Müller biogeographical appears toprefer algae brown that sectors with infected by has been frequentlyof observedaggregates inthe Adriaticcells inwhich the alongzones thef fragmentation of filaments of the progressivegrowth of theaggregates dimensionalintheir propagation inorganic inside.Asreported debris contained int grayish ofpigmentscolor algal loss and causedthelargeby the cells amount the oforganic in and the reproductive structuresareabsent,aggregates and compact anddisplay thebecomemore a attached butduring of species, dimensional thei this lose growth, form their the filaments polarity cellswelloccur andstructures pigmented andreproductive they because inthe typical Algal DISCUSSION fattyacids ratios C15br/C15:0 than themucilage andhigher produced by Inmucilage. contrast, ratios of and polyunsaturated acids, fatty diatom ratio respectively.and The C16:1/C16:0 andC16P/C18P marina both the algabenthic andFAs mucilageby mucilage. its majorin produced The ofbacterialHowever, the contribution fatty andthe acids ratio were C15br/C15:0 in quite similar (preferablygreen) or (diatoms) microalgal sp diatom andC16:1/C16:0 crinita

Acceptedconenoses. Article ) is that its mucilage) isthatits

(Banjo N. marina N. Eurychasma dicksonii Eurychasma le Island)wereC18:3(n and le C18:1, C16:0,

ilaments andinthe apicalilaments pseudo In stag thefirst

mucilage respective were tothe ratios also obtaine similar N. marina N. ratios (Table 4), thus contained a measurable proportion of C16 PUFA in addition tohighera proportioncontained inaddition of measurable PUFA C16

es of their development, thefilaments theirof development, es of mucilage acontained proportion significantly higherofbacterial

A. crinita

(Wright) a parasite Magnus, largelyspr , which cancaused be by abscission by senescence, ecies tothe total organicecies matter ofaggregates.

revealing the contribution of other macroalgal contribution of revealing the he previous - hairs, latter aspect this and infection; by fungal

- 3) among thesaturated, monounsaturated, paper (Sartoni et (Sartoni paper al.2008), filamentous habitus(Konno andfilamentous becomes A. A. A. crinita A. ead invariousead crinita A. crinita d for

possible throughpossible the Nematochrysopsis Nematochrysopsis

A. crinita

(Table 4). show an evident

often are r polarity, the

This articleThis protectedis by copyright. All rights reserved. greater diffusion resistances, fresh therebysupplyingcell and at with C promoting higher the rates referencesgreater The therein). turbulence maintainslower towhichplanktonic algae are exposed community metabolism th Negro et al.2005). higherDOP,concentration which could of hydrolyzed bepartially phosphatasealkaline (Del by to the observations ofMisic et al.(2011). regeneration The pre ofwas P 2005 prokaryotic activity insidethese aggre findings reportedet et by Misic and al.(2005). al.(2011)could Giani Thissituation bedue to of incidence inthe in DCHO) have matterfound and (DOC ofdissolved organic been regenerationNutrient andorganic mattercycling temperature thefavor trend spread of could Northernreported Sea Adriatic as which agreement isin theincrease with observedfluxes inair and temperature heatinthe annual 1986 however, t Inof Mediterraneanwaters. exceptional warming previouswas 2004,no studies; reported by mucilage formation, i.e., 2010), information lacking. onthe aggregates filamentous formedby these algae is the benthic respect aggregateswith thatinthe surrounding substantiates to which previous water, The boundarydevelops inregulating that layer over algae benthic mightafactor key be benthic The DIN/PO According etal.of (2007), macroalgal inbenthic thespread species toSchiaparelli involved Although , -

Accepted Article2005 showed a significant Misic etMisic al.2011). 13 C he analysis performedheal. by Tedesco(2007) et for thesurface water - depletion (France 1995).

A. crinita 4

ratio was the higheraggregatesand outsideof decreased markedlyinside,similar

has been reported in different sub has reported been indifferent

A. crinita, rough ofnutrients andcarbon recycling internal (France 1995and

increase in May, June and JulyincreaseJune in May, Adriatic study inthe north sites, by Russo et al. (2002). Therefore, etal.by Russo (2002).

might by befavored in the

gatesenzymatic canbe where high activities ( A. crinita A. . Higher nutrientand therelevant concentrations

at higher latitudes - basins ofthe Sea Adriatic ( creasing ofthe temperatures surface we hypothesize thatth we hypothesize . sumably favored by the

temperature during

terstitial waters Del et Negro al. Antolić et al. Antolić et

is even even This articleThis protectedis by copyright. All rights reserved. andmucilages involve aging selectiondevelopment of morea from diversecommunity tothe composition partially reproducesresidentthe microphytobenthic diatoms the community, occurring of aggregates. inthe development contribution respect ofdiatomswith phytoplanktongroups theotherthe metabolic to pathways to from previous observat constituent inmostcommonlyfound Tyrrhenianbenthicaggregates,the water column when hypothesis byet Misic raised al organic2005). ofnutrients Moreover, the matteraggregates and enrichment insidethe supportsthe population, aspreviously showninthe studiesperformed inthe and leaves heavier marina δ bacterialelab associatedmechanisms with differentthebenthic thatoccurregeneration layer, nutrient in and i.e., NO wateralgae.Furt of and waters inthe the surrounding of DIN (nitrification/denitrification etinthe processes; interstitial Cole al.2004) thatoccur inside experienced i.e.,nitrogen irradianceand high fixation naturalNsources, aggregates of carbohy Istrasurveysattributed topreferentialbacterial in couldduring aging be degradation ofthe 15 N inalgae in inparticular Diatom communities establishedDiatom in the aggregateswereAdriatic microphytobenthic the Nutrient recycling in The This situation The 3

Accepted Articleor NH

13  (Tablecould 2),bethe result ofbacteriaactivity thatuses pre 15 C depletion theC with C N 4

values found inthe benthicbe theresult algaecould ofdifferentfractionation processes by plantsand oration. High concentrationoration. of the bothnutrients High aggregates, inside withmore positive

can be typicalgrowing ofmacroalgaesystems oligotrophic inshallow that have 15 they areinducedthey by currentswindmixing. dispersed N isotopes inthe substrate (Dai etal. 2005 drates, which tend tobe drates, ions the aggregates thethe candevelopmentan phytoplankton abundant support of algaeet al.2013). (Raimonet The (Innamorati A. crinita . org

(2007)aggregatescan that asa theact so /N increase observed inbenthicalgaecollected during observed successive/N increase

sampled in Tremiti and Istrasampled inTremitiand and inJune) (sampled et al et

From the earlierstagesthe in which From micro .

2001 13 C enrichedal.(van 2002). et Dongen ,

Sartoni et al.2008), confirming thus themain

her during fractionation occurs of assimilation

increase of increase of ).

Tyrrhenian SeaTyrrhenian (Deetal. Philippis

ferentially light δ (Lamb etal. 2012 15 urce for ofnutrients the as previouslyevidenced N values could be

algae

14 N isotopes N. N. ) .

This articleThis protectedis by copyright. All rights reserved. heavilythe most samplesfromand bothpelagic benthic aggregates degraded mucilage showed galactose2005) witha culturesin axenic by( species diatom several lowest galactosecontents (Giani et results exopolymers al.2005).for Similar were produced found reported, exopolymers produced freshly et al.2005, stage mucilageaggregates, bothmarine culture of asaxenic for (Giani previouslyand used samples observed resembleaverage ofthe high the composition approximately (Sartonisuggesting thus 11%Adri et ofxylose al.2008), thatthe polymerchain values Mw=800 exhibited of (about kDa) approximatelygalactose w/wof 36% and polysac afterfractionation, polymer as (Sartoni previouslyetlong reported al.2008). Those notably for tothat thehigh similar found lower vs. content18%) (5% of xylose with the Tyrrhenianaggregates percentage displayinggalactose a(50% higher and vs.30%) of galactose respect with Tyrrhenian contents and the xylose to produced by sample 2008) in isshown sites)with thoseaggregates obtained (Istra and from Tremiti theTyrrhenian Polysaccharidic matrix phosphorus ( organic and2008)etal. (Sartoni likely able are to exploit availabilityhigh the internalof rapid recyclingof and abundantrecurrent inboth pelagicdiatoms ( monospecific nearly dominanceof AcceptedUsinggalactose thesumofdeoxysugar (fuc+rha) contents and of as anindex Article charidesby formed approximately 3500 in Sartoni et al. 2008 Sartoni et al. Fig. 6

Fig might Najdek et al.2002). ure

content lessthan 25% (fuc+rha) sum content andgreater a than 45%. Conversely, .

A comparisonaverage the of of monosaccharide patterns Adriatic

be ascribed todifferent degradation stages. 6 . The average Adriatic from samples values , Urbani certain etal.otherbe featurescan As 2005,), highlighted. Cylindrothecaclosterium. . The average composition found for Adriaticaverage foundfor composition samples. The is

andsum (fuc+rha) aggregatesshowedand thehighest -

molecular

Magaletti et al.2004 Magaletti - molecular - 4000 monosaccharide connected4000 residues inthe Najdek et al.2002 - weight fraction of weight fraction

- weight and exopolymers, thedifference

This species This i , Myklestadet 1974,,Urbani al.

) and benthicaggregates )and A. crinita differ significantlydiffer for both s one ofone s mostthe Sea (Sartoni et al.

atic samplesmay

the degradation exopolysaccharide A. crinita - chain

algae

This articleThis protectedis by copyright. All rights reserved. marina or larger, clouds (14.2 i.e., mucilaginous aggregates(Najdekal.were 2002)andeither et smaller, i.e., ratioscontribution were andC15:br/C15:0 (0.9 muchlower Fatty aci Fig. 7). by 2008). Thislarge polysaccharidefraction again represents produced thefreshly exopolysaccharide Fig.(point 7in exclus from 7)obtained sample size 5using Adriaticin Fig. comparison sampleinaddition to (point3the of 7) highfraction molecularweight from 5 and 7fromsamples). the Tyrrhenian As observedfor produced by the Adriatic aggregates. polysaccharides activity of adepolymerization with extent bacteria lesser tothecolonize due that fraction (point6in Tyrrhenian (point 4inFig. butare sample molecular benthic high 7) toits quite close weight the Adriatic 1 samples (symbols previous (Sartoni paper al.2008), et Sartoni et al.2008), revealing thus and specific enzymatic preferential degradative processes. markedfucose and decreases andgalactose rhamnose of an of increase InFig ComparingIstraTremiti sampleswith theTyrrhenian the and N. marina, N.

Accepted 5inFig.galactose a withrespectthe Tyrrhenian higher sea(point content 7)showed the of to Article

(24.9%; shownin 12.6),as d signature ure

7, the averagedata fo7, the Nematochrysopsis marina Nematochrysopsis

which is quite similar ingalactose similar tothat foundinthewhich Adriatic content isquite sea(point3in Fig. 7 . In benthicbyformed mucilage the

). Thi - 17.1%; 5.5 s observation suggests freshs observation suggests for algal the production Adriatic and 2inFig.arethe (unfractionated) 7) from different that of r the monosaccharidic composition ofexopolysaccharides composition r the monosaccharidic Fig it isnoteworthyit (fuc+rha)galactoseand that the percentages of - ure 6.0), particularly from6.0), formedparticularly benthicmucilage by

are also reported(point3from samplesand Adriatic points

8 .

A. crinita A. crinita, A. - ion chromatography(Sartoni et al. 5.8%; 0.98) 0.0to thaninthepelagic A. crinita A. , the the bacterial fatty acid N. marina N. residues (Giani et al.2005, residues

flocs (3.1 flocs sample reported ina

exopolysaccharides - 6.8%; 2.5 N. N.

- 3.2) This articleThis protectedis by copyright. All rights reserved. all benthic mucilage investigated inthis abilityNajdek toreproduce etTherefore, (Grossart 1999, and inmucilage al.2005). incontrast with or as selected species,i.e., polysaccharideproducers ( a matrix of mucilage I closterium bacterial observedcontribution was pelagicaggregates inaged dominated bydiatom advanced tookplacethe mucous degradation matrix of intheseaggregates. benthic Such a high benthic by mucilage formed were growthfoundof orinhibit torepelpropagules microorganisms. 1982), sesquiterpenoid brown algae, diterpene and alcohols etal.Towers i.e., (Schmitt 1995),brominated (Phillips phenols secreting mucus( defensivecompounds ( marine algaeefficient eitherevolved have epibiosis, chemically tocontrol strategies by producing and freshand notconsiderablycolonizedby thus bacteria thetime ofsampling. during Many mechanism in Thesegrowthconditions for might bacterialeither more indicate a unfavorable efficient highly contrast, thebacterial coloniz aggregate self In interact process,andform the bacteria this of phytoplankton pelagic the foundation with contribution increase theaggregate process, parallels aging i.e., usuallyIn foundpelagic within mucilage. thepelagicmucilaginous the aggregates, bacterial es n pelagic aggregates, the most prominent diatoms contributionsof were most the aggregates, n pelagic as either documented tablished within anythesampledtablished within benthicformed mucilage of by Accepted Article This observation suggested

( Blažina et al.Blažina 2005 et - A. crinita sustainability throughsustainability theproduction Steinberg etSteinberg al.1997 s (de Nys et al. 1998), and halogenated Nysands (de etal.(Steinberg furanones 1998), et al.1997), Nylund and Pavia 2003

for from extrication orthat epibionts the mucuswas continuously exu C. closterium,C. N. marina N. ation of thebenthic mucilage formed by

, that conditions for heterotrophic bacterial growth werethat conditions forgrowth heterotrophic bacterial not Najdek et al.2005). ,

was heavily colonized by bacteria, thus suggestwascolonized by bacteria, heavily thus Nylund and Pavia 2005 Nylund and study

that can survivethat can such ( , Fig. 9 Fig. Fusetani 2004 Kaltenböck and 1992 Herndl -

decomposition cycle (Najdek et al.2002).

much higher ) a ). Antifoulingred metabolites of and extreme environmentsandhave the degradation of the mucous matrix. of themucousdegradation matrix. nd/or physically and by sloughing A. crinita A.

values ofthevalues ratios I A. crinita A. n contrast with n contrastwith

with respect those to , Degobbis et al.1995

was negligible Cylindrotheca A. crinita ing that ing ded . In , ) This articleThis protectedis by copyright. All rights reserved. degradation organic constituents. of the aggregates suggeststhattheexoenzymatic results inanon activity of these the aggregates, whereas necessary toverify conclusio this moreto its recalcitrant ortoalgal nature intensive studies are defensemechanisms,althoughmore forrecognized bacterialgrowthconditions unfavorable inbenthicmucilagemight beascribed either mucilage, formed thebacterial by contribution constituent andexopolymer diato increasing water trend insurface temperatures. The spread of CONCLUSIONS as a mucous matrix, mucilage formed by from todominant contributions(diatomratio: low 0.5 contribution ofdiatomstothefattyacid compared pool ratherof phytoplankton inthe mucilaginous thanactivegrowth. occurred matrix The relative (0.0combined withthe lowC16P/C18 phytoplanktongrowthall ( benthic mucilageto theC16:1/C16:0 large in due of variations presencegrowing ofactively mucilage inpelagic diatoms (NajdekThe et conditions al.2002). for (0.6 C16:1/C16:0 the aggregates and in However,nutrie enrichment of

Acceptedms, thefattyacid monosaccharide and indicated signatures that Article A. crinita

- 1.1) and C16 lso observed in the microscopic inspection.lso observed inthe microscopic N. marina N.

in the Adriaticalso thenorthernbe and in favored could by region the the surrounding waters of the algae surrounding of waters the - producing species in thebulkaggregate.Inproducing in pelagic species contrastwith



15 P/C18P (1.3 suggested considerably not did contribute thatdiatoms tothe N values depended onthefractionation processes thatoccur inside n. Theof isotoperatios nts and dissolved organic watersnts anddissolved matterof the inthe interstitial - 0.4) ratiosand indicated different that occasional embedding

- 6.2) as well6.2) as the

A. crinita A. Although the benthic aggregates are aggregates colonizedby thebenthic Although

- 1.7). obtainedThe for results similar benthic with that of other microalgal classeswith that ofother varied

δ was lowin quite 13 . C

demonstratedthe marine origin of diatom ratio (0.5 diatom ratio

- negligible the contribution to A. crinita A. benthic mucilage. The

was themain - 1.7) suggested the suggested the 0.1 - 1.2) This articleThis protectedis by copyright. All rights reserved. Calvo Burney Bradford Blakeney Athanasiadis Aspila Antolić Aktan environment. tobothMediterranean, understandexists their theneedecological impact and onthe their role same algalgeographical ofthe and originirrespective site marine physicochemical features. analysis composition pattern, suggesting thus aggregates produced thesame thatthese have the Because alga differentbenthic Compared with Tyrrhenian benthic aggregates produ coasts during 1991 procedurecarbohydrate for total analysis and polysaccharideestimation of proteinof protein using theprincipal acetatesalditol monosaccharide for analysis. (Rhodophyta,Ceramiales). inorganic, phosphate insediments. organic and total Heterokontophyta: Phaeophyceae. 2010 17 benthic mucilaginous aggregates Sea inthe Aegean (Eastern Mediterranean). ,

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142 33 . , Ehrhardt . -

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K - . . herbivore interactions: I. interactions: herbivore

, Ehrhardt , 125

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This articleThis protectedis by copyright. All rights reserved. Nikolić Naj Najdek Najdek Myklestad Müller M Moncreiff,& M.2001. Sullivan, C.A. & M. Mistri, Cec ü ller, D.G., Kapp, M. dek the Adriatic Sea. Acceptedinvasive red alga intrusions. closterium Res different of compositions types aggregatesAdriatic. ofmucilaginous inthe northern during the1991 nine differentculture. species in eukaryoticalgae(Phaeophyceae). brown parasites inmarine ArticleEurychasma dicksonii 49 106 seagrasses stable from multiple isotopesanalyses. beds: evidence 63 Paramuricea clavata - : ,

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Unusual changes of zooplankton fatty changes zooplankton Unusual acid inthe northern of composition Adriatic .

mucilage event.mucilage , P ,

Acta Adriat Acta . Womersleyella setacea

M ,

A roduction of carbohydrates by marine planktonic diatoms I of carbohydratesroduction by planktonicdiatoms marine Comparison of ,

F D . . . . & , Đakovac

, Antol , . I . C Res . , Mioković

(Oomycota) and . & Knippers -

Short termShort impacts atthe populationandcolony levels. .

27 Küpper ić : . 851 ,

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. Ecol 1999

Chytridium polysiphoniae Chytridium Ivančić - Mar

202 (Hollenberg) (Rhodophyta, R.E.Norris Ceramiales) in .

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, Balata ,

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P . Invasions ofalien macroalgaeMediterranean in , Nuccio – . . , Sartoni

, 198 299 .

F [E 143 Decadal Climat . lgol A .

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22 111 . . :

] 875 1963 . , , :

209 28 The Sea

C Bot , J. FeldmannJ. e –

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. & 882 . - . 289 Mar

218 The influence oforganisms onthe Mar

Giani .

Cattaneo . -

. . .

301

23:23 Vol. 2, Wiley, LondonVol. 2,Wiley, . Ecol

ic Anomalies inthe Northern Adriatic (Sarcinochrysidales, Chrysophyceae), 38 ,

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: 121 .

. - 23 Acinetospora crinita Acinetospora

30. 2008 - Vietti - : 12 Acinetospora crinita Acinetospora 340

e osservatesulle coste 5 . . Benthic mucilaginousBenthic – ,

351 R .

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This articleThis protectedis by copyright. All rights reserved. Vizzini van Dongen Urbani Tsiamis Tedesco Steneck Steinberg Socal Schmitt costatum marine diatoms coasts. biogeochemicalyears variabilityin the last20 functional groupapproach. microbial colonization. features 2003 Adriatic of the Sea Northern inthe period Lazzari functionsMultiple for seaweed metabolites. secondary coastal lagoon. and(13C/12C 15N/14N) ofprimary consumersMediterraneanand ina producerswestern 232 monosaccharides aquatic andof lipids plants. algae and terrestrial

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G , ,

, , ,

R : ,

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T R K , Acri , L . . &

, Magaletti . P . . . I. Phaeophyce , Hay , – S , Panayotidis , Socal

, . A 92

. , Schneider . &

B Mazzola Effect ofP . , Bandelj, , .

F ,

. Watling . M , Schouten , Bastianini ,

G Cylindrotheca pseudonana closterium, Thalassiosira Mar & . , .

, Bianchi , E

A Lindquist , . , . ,

, Sist , -

V ae. ae. , R . Biol. depletion andgrdepletion P

L 2003 . . & . Biodegradation , Cossarini , , Economou . ,

Mediter.

1982 , S ,

Kjelleberg M P . & 142:1009 , Mar .

. & F Seasonal thestable in nitrogen variations carbonand isotoperatios . , , Bernardi Aubry .

. , N Damsté

Feeding capabilities and limitations ofherbivorousFeeding andcapabilities limitations a molluscs: . Acri Cicero

. Biol

1995 Mar ,

G - , – Amilli

. , . & F

1018 owth status. owth

. 68 , S . .

, Veneri , Sci

J 8 . Constraints onchemically A

. : : Solidoro 1997 S 299 211 . . M . .

, S 14

A . . -

–

2002 2005 319 : . . (1986 , 220

141

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D .

, . , Bianchi Katsaros

Sci . . - . . C

147

& Carbon isotope variability in Extracellular carbohydrates – .

2005). .

2008

Vichi

Total Ecology . –

2006 , , .

, . C F Hydrological biogeochemical and

Biogeosciences

M Environ . . . , Cassin

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449 : Prog 673 –

306 released by the – – 468 . , 687 .

Ser J . , De

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This articleThis protectedis by copyright. All rights reserved. external the aggregateof: to a) phosphates Fig. water inthe interstitial 5:Concentration thebenthic to internal and aggregates inthe seawater unpublished data, and (****) Giani et al., 2012. aggregates sampled Istra. in aggregates of A. crinita pelagic mucilages (**) and phytoplankton (***); Fig. 4: Croce shallows, and Porer within an aggregate the in early stage development of (Tremiti Islands, bar=500 µm). Sauvageau within an aggregate (Tremiti Islands, bar=500 µm) growing onthalii of Tremiti Islands. Fig. Sampling 1: sites: a) in northern the Adriatic Sea, Istra peninsula; Welker White Fig. a) 2: Fig. 3 128 Harvey)ex (Ectocarpales, Sauvageau Phaeophyta)nell’Alto Adriatico. detrital microflora. biochemicalthe determination for methods of mass andcommunity fungal structureinestuarine

Accepted Article , : a

: ) C ) Relative abundancesRelative ofthe dominant microalgaeaggregates inthe benthic (Secca of Rovinj

D 827 C , phytoplankton (***), and pelagic mucilages (**); Aggregate on . . & C org - . /S vs./S C , Bobbie 829 A. crinita

Bressan

Dictyota org ,

/P /P

Cystoseira corniculata , sampled the in Istra and Tremiti islands. The linear isfit shown for the R

Bot G . A. crinita A. J .

. Data from (*) 1994 , Nickels .

Banjole Island) and Tremiti IslandBanjoleIsland) and Tremiti inMay (Ferraio) sampled 2004. sp. (Tremiti Islands); c) filaments of

Mar . . Aggregati mucillaginosi bentonici di

aggregates from the Adriatic sea and Tyrrhenian sea (*), in 23 ,

: J 239 . S

Sartoni et al. 2008,(**) Giani et al. 2005, (***) Berto, . , Fazio – 250

(Turner) Zanardini (Rovinj, bar=1 cm; b) (PO

b .

) δ ) ,

4 S ), silicates (H 13 . D C andδ . &

c Davis ) δ ) ; d) d) ; apical portions of 15 N different in benthic agg 13 C vs. C Acinetospora crinita ,

4 W SiO

b) in the southern Adriatic Sea, . M 4 org . ), b) nitrates (NO

1980 /N Acinetospora tot

in different benthicin .

Giorn. Nonselective Acinetospora crinita

(Carmichael)

small aggregates Bot regates ofregates

crinita 3 . ) and

Ital

. (Carm.

This articleThis protectedis by copyright. All rights reserved. pelagic (flocs and clouds, Fig. 9:C16:1/C16:0 marina Fig. 8 obtained from paper the previous et (Sartoni al.2008). high weight molecular fractionsofsamples 5),re 4)and Tremiti, 3) shown Nematochrysopsis marina Islands.valuesForcomparison, from theaggregatesof Fig. 6 and d)organic dissolved carbon carbohydrates(DCHO). and (DOC) dissolved (NHammonium Fig. 7 : Accepted: Article :

Relative monosaccharide compos Relative monosaccharide

, Bacterially fattyacids derived vs Galactose percentage ondependence ofrhamnoseIstra, and percentage fucose for 1) 2)

) and pelagicmucilages (flocs, N. marina N. 4 ), c) dissolved organic(DON) nitrogenand phosphorus organic dissolved (DOP), vs , Istra,, 4)

C16P/C18P ratios thebenthic in C16P/C18P (

sampledIsland at (Istra) Banjole and inthe Tyrrhenian Sea are also ) mucilages. A. crinita, .

ition of ition

C15:br/C15:0 ratiosthebenthic in C15:br/C15:0 (

Tyrrhenian Sea,Tyrrhenian 5) a nd clouds, nd A. crinita

A. crinita spectively. Data of Tyrrhenian Dataspectively. are of samples

A. crinita

aggregates from the Istraaggregatesthe from and Tremiti ).

N. marina N.

sampled inthe Tyrrhenian and Sea , , and , Tyrrhenian Sea, 6) and 7) 6)and, Tyrrhenian Sea, 7) N. marina N. A. crinita

, ,

) and and N. N. This articleThis protectedis by copyright. All rights reserved. Istra N. m Istra Istra i Tremit Istra crinita A. reported. Table Elemental2: composition of the benthic aggregates. Averages and standard deviation DiavoloPunta Secca Vedove Ferraio Cretaccio Tremiti Islands CrocePunta Porer Banjole Istra Table 1

Site

arina

Accepted Article Site

Sur

26/06/2004 26/06/2004 29/05/2004 26/05/2004

3/05/2004

face the sampling bottom and depths near date

26/05/2004 25/05/2004 26/05/2004 25/05/2004 25 29/05/2004 05/02/2004 26/06/2004 30/05/2004 05/02/2004 25/06/2004 29/05/2004 05/03/2004

/06/2004

Date 12 12 16 17 Dept

12 m h - - - -

16 15 18 24

Depth /m 89.63 90.22 91.55 88.53 5.09 2.85 H 90.46 7.7 % 2

1 1 1 1 1 1 1 1 1 1 1 1 1 0

22.65 18.75 25.71 30.78 32.52 4.17 4.11 4.56 C Depth /m % tot

17 16 16 16 16 16 16 15 15 15 24 18 22

C 12.87

4.38 4.97 2.35 5.85 1.56 4.67 4.82

inorg %

18.28 13.78 19.86 26.11 19.65 C 4.97 2.56 5.5 % org

0.79 1.05 0.23 0.49 0.22 2.22 1.7 2.3 N %

nd 0.58 0.04 0.63 0.12 0.68 nd nd % S

s (in italics) are

nd mg 571.22 456.74 768.37 209.28 778.98 501.77 80.26

P . kg

26/06/2004 26/06/2004 30/05/2004 29/05/2004 29/05/2004 26/05/2004 26/05/2004 05/02/2004 05/02/2004 Date This articleThis protectedis by copyright. All rights reserved. Adriatic aggregates. Table 3:Monosaccharide(% composition exopolysaccharide w/w) of thebenthic in fractions Istra N. marina Istra Istra i Tremit Istra crinita A.

Site

Accepted Article

Diavolo Banjole Banjo Banjole Banjole Station 26/06/2004 26/06/2004 29/05/2004 26/05/2004 Ferraio 3/05/2004

Croce Punta Punta Punta Porer Porer date 2

le -

Nematochrysopsis

Acinetospora Acinetospora Acinetospora Acinetospora Acinetospora Acinetospora Acinetospora Acinetospora Depth 12 12 16 17 12 m Algae - - - -

16 15 18 24

- - - - -  22.78 20.92 20.88 18.03 18.91 0.71 0.73 0.69 %o 13

Depth 15m 15m 15m 16m 16m 22m 15m  4.56 5.29 0.44 3.06 0.59 5.17 2.68 1.3 %o

41.5 39.3 29.3 28.7 32. 31.8 32.5 Gal nd mole·mole 35 27 C

1033 org 622 524 871 164 896 362

/P 14.8 12.6 14.4 Glc

5.7 6.6 4.1 3.2 tot 32

nd mole·mole Man 10.9 14.3 10.1 12.9 11.4 10.3 N 9.7 6.9

9 40.7 39.9 65.9 13.8 99.3 tot 4.9 65

tot

11.5 29.2 26.1 21.3 40.4 13.8

Xyl - 7.6 1 11 17

mole·mole

C 26.95 15.3 13.5 13.5 10.2 Rha 2.2 2.2 org 6.9 9.6 5.6 6.7 5.4 3.2 7.9 8.4 2 3

- 1

12.2 20.2 11.5 11.1 20.8 17.9 28.6 Fuc 4.1 4.9 nd mole·mole

63.2 16.4 90.7 26.4 83.8 org nd nd Rib

1.5 0.7 /S 2 0 0 0 0 0 0

- 1

22.3 Ara 1.2 2.8 2.9 2.9 1.2 2.1 0 0

This articleThis protectedis by copyright. All rights reserved. Accepted Article traces; na 18P 16P, /C16:0); C15anteiso + + +C17iso C17anteiso C17:0 +ratio diatom C18:1n7); of the isolated al Table 4 : Fatty acids Sample DIATOM RATIO C16P/C18P C16:1/C16:0 C15:0br/C15:0 BACTERIAL C22:0 C22:1 C22:6(n C20:0 C20:5(n C20:4(n C18:0 C18:1(n C18:3(n C18:2(n C18:4(n C18:3(n C17:0 C17:0br C16:0 C16:1(n C16:3(n C16:4(n C15:0 C15:0br C14:0 C14:0br

Fatty acid composition ofthe benthicFatty mucilage composition acid produced by –

not applicable not

------ga 3) 3) 6) 9) 3) 6) 3) 6) 7) 3) 3) –

A. crinita A.

sum of polyunsaturatedres acids fatty 16and18Catoms, with

. br 34.15 ±8.71 16.19 ±9.09 14.82 ±4.41 0.51 ±0.34 0.95 ±0.82 0.04 ±0.13 0.57 ±0.51 3.49 ±2.35 2.26 ±1.46 2.76 ±1.66 7.14 ±2.86 5.06 ±2.89 4.36 ±2.33 3.01 ±2.49 0.58 ±0.50 0.62 ±1.19 0.12 ±0.34 0.49 ±0.45 0.77 ±0.81 1.31 ±0.78 0.24 ±0.33 1.02 ±0.41 0.13 ±0.15 0.53 ±0.35 0.25 ±0.36 0.36 ±0.52 BENTHIC MUCILAGE A. crinita –

branchedacids; bacterialfatty fatty acids -

N. marina N. 12.59 14.32 12. 24.87 10.72 23.51 4.13 1.14 3.96 4.16 0.88 0.32 0.54 0.34 1.39 0.53 5.98 5.61 5.58 0.82 1. 1.90 2.26 tr tr tr tr 07 59

14.50 ±0.49 41.22 ±1.16 7.96 ±0.56 0.92 ±0.35 0.24 ±0.32 0.00 ± 0.19 ±0.02 0.20 ±0.28 1.69 ±0.18 0.93 ±1.31 0.40 ±0.57 0.18 ±0.25 1.08 ±0.01 2.62 ±0.47 2.48 ±0.31 2.28 ±0.08 9.84 ±0.61 4.76 ±0.23 5.35 ±0.19 2.90 ±0.08 0.28 ±0.39 0.35 ±0.50 1.45 ±0.51 A. crinita ALGA na tr tr -

A. crinita –

(C14:0 + C16:1 + C16P +(C14:0 +C16:1 C16P

–

(C15:0 + C15iso +(C15:0 +C15iso

and pectively; tr N. marina N. and -