This article isThis article by protected copyright. All rights reserved. 10.1111/jpy.12614 doi: d to lead copyediting, paginationbeen throughthe andproofreadingtypesetting, process,may which This article acceptedhas been for publication andundergone fullpeer review buthasnot LaboratoryEnvironmental ofabbreviations: List Pterosiphonia Key Running * Vancouver, ColumbiaBritish V6T 1Z4 ofUniversity Columbia, British andBiodiversity BotanyDepartment Patric ofUniversity Washington, Department ofBiology, Seattle, Washington 98105 ofUniversity Washington,Harbor Laboratories, Friday Washington Harbor, 98250 Emily ofUniversity Washington,Harbor Laboratories, Friday Washington Harbor, 98250 Kevin Miklasz ofUniversity Washington,Harbor Laboratories, Friday Washington Harbor, 98250 Vancouver, ColumbiaBritish V6T 1Z4 ofUniversity Columbia, British andBiodiversity BotanyDepartment Rebecca Guenther Macroalgal sporedysfunction: : Article type Letter 0000 ID: (Orcid GUENTHER REBECCADR. Accepted GuentherCorresponding Author:Rebecca ( Article words: algae,words: k

Carring T. ifferences between the of thisversion citethis article and asVersion Record. Please title Martone : Ocean acidification: Ocean impacts adhesion spore , ton

Polyostea

climate cha *

FHL, Laboratories; Harbor OAEL, Friday Ocean Acidification

, pH

nge, o adhesion, cean delays acidification andweakensadhesion -

0002 - life cycle, 3489 [email protected] - 0561) propagules,

shear stress, shear stress, Research Centre, Research Centre, )

Corallina

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This article isThis article by protected copyright. All rights reserved. forces imposedhydrodynamic waves, bybreaking withenvironmental stresses, butalso such etTaylor al. 2010 perturbation attachment onspore understood see ispoorly (but Steinhoff 2011, etal. t spore conditions, r differentially verification isneeded. via and glycoproteins) protonation displacement, andcation although properdisrupts curing of andgelformation spore adhesives(anionic polys strength at dislodge lower flow by 40 non rhodophyteintertidal macroalgae, on flu impact biomechanical criticalthis adheresettle, andcontinuethealgalcycle life acidification he successfulsettlement andattachment ( spores ofalgal

Accepted Article me to examinetheeffect of - calcified ( - dys 52% The establishment ofmacroalgal andpersistence benthic dependupon communities E arly life stages lifestages arly of function, regardless of the tolerance physiological mature of thalli.

P. and life stage life . educed , F

robusta impact Polyostea robusta or macroalgae

weakened ). In the intertidal zone spores ofrockyshores, must notonly with contend

ocean Padilla s has been . -

Our of reduced of reduced pH Results are are Results consistent with thatreducedpH ourprediction induced shear forces, induced shearforces, spore adhesion of marineare organisms attachment pH - r Gamiño 2016 al. et esults demonstrate , reproduction and reduced has the potential to has thepotential largely overlooked ). Red uced pH delayed spore uced pHdelayed e calci on

in two macroalga strength pH early on fied ( but had no effect but hadnoeffect , yet spore ) spore adhesion , yet , yet impactthe ofenvironmental

in that oceanthat

predicted to population persistence C .

C. vancouveriensis impact macroalgal communities via the effect of oceanthe effectof acid W orallina attachment e explicitly tested e explicitlytested the

Chamberlain 1976 e. If hold results

acidification negatively vancouveriensis Fletcher Fletcher attachment . W be o

n time

vulnerable vulnerable e the attachment developed and , and causing spores to sporesto causing experimental experimental

, Call rely onsporesrely to in field in field Santelices 1990, of both of both ification on strength ) andone accharides to ocean to ocean ow 1992 a

shear

species

, in and ,

two

This article isThis article by protected copyright. All rights reserved. pH reduced often understoodand are poorly 1972),and Jones production and chemicalthe composition (Boney1981,Fletcher and Callow 1992), curingrate (Moorjani stiffer, more compact be whichmay also gels, less deformable(Picone conditions, gel causingpolymersand potentially and ( tocollapse entangle weakeni anionic and Mooney2012,Li spore general, c 1976) strength ofthis securing spore ( dischargespores asecond mucilage, ofadhesive type composed mainly glycoproteins of the 1981 polysaccharides ( initially released boundarylayer the to avoid being algal (andthus algal spores thalli)dependsuponthe s as increased temperature Accepted ArticleChamberlain a strength , Fletcher andCallow, Fletcher 1992

, likely increasing ioniccross dueto mucilages,

Settled polysaccharides to become protonated ng hydrogen bonds wit ross ,

of mucilage this determines time attachment ( we hypothesized sporeadhesionmaywe hypothesized that be -

s linking is

spores viscosity 1979 nd Evans1973,Pueschel secondary mucilage to substratethe ( from sporangia and shiftsocean in may ionic pHlinkages (Verdugo affect Chamberlain andEvans 1973, must et et al required for required the proper , generally increases generally desiccation .

resist shear forces shear imposedresist velocity agradient in water within by 2013). h water, , given ). This mucilage). This initially adheres sporestothesubstrate, and

Chamberlain 1976, enveloped bymucilage composedofacidicsulphated rate dislodged ( R

increases through time educed pH educed

(Gunn and the displacing , -

perhaps linkages oftheadhesive( generalized response et al.et ( Bu Vogel 2003 )

(Lee Mooney and 2012,Li , which

can carboxylate cause and groups sulphate on formation of Boney 1975 et al. et

1984) of spore mucilages1984) ofspore , stabilizing

seawater pH.seawater survival Thus, initial of Fletcher Fletcher and peed andstrength of

2005, spreads outform to a pad,permanently ). To inattachment, aid

affected affected Boney

Bu al et (Charters (Charters , Lee andMooney2012 marine Chamberlain 1976, cations (e.g., Ca of anionic hydrogelsof anionic

1975, Callow Callow 1992 by ocean acidification. .

Jones etal. 2005) hydrogels, hydrogels, et al.et

et et al. 1981 .

et al. et spore spore 1973, Chambe In low pHIn low vary

et al. et ). Once attached, ). Onceattached, 2011). 2+ ) .

including 2013) , 1982). The adhesive

attachment. across taxa K s Boney 2004, Lee pores are pores + ) , Although ,

Na

making , to In + )

rlain rlain ,

This article isThis article by protected copyright. All rights reserved. to each Prior shear flume test, the on fitted the was bygenerated water each column calculated were accordingto Schultz height column wasvaried arangeof height tocreate flume was allowed toset. in resulted a of gradation attachment time theworking across section attachment landingend oftheplate.Spores inthe section working water hr documented nearshore environments in inWashington ( seasonallyrange from to8.0(Mur 7.6 InformationSupporting water carbonate chemistry analys CO bybubbling were established of pH andtemper OceanAcidificationthe Environmental Laboratory FHL, (OAEL) at allowing precise control toexperimentalexposed conditions while releasing s testing. HarborFriday maintained Laboratories (FHL),and anoutdoorseawatertable in WAIsland, bipinnata -

Accepted1 Article ) drove ) drove reproductive algal thall , while r

release

The spore settlement The spore ofacarriagesystem consisted apparatus specimensSporophytic of Reproductive parent a frondswereheldat ) and

time, determined by the ratetime, parentthalli bythe weredriven determined the across plate (Deadman 48°30'48.93" Bay: d eleasing spores (Fig. 1a sporeseleasing (Fig.

Corallina vancouveriensis a tall columna tall thatflushes quickly ofwater acrossthe ature ature in aflow After sporerelease andsettlement, a ). Ambient fortheS pHvalues - through system2013 etal. (O’Donnell 2 es macroalgae

i across glass a plate (0.6 x 7.6x60cm) (7.75, according to Dickson( ). Theworkingsection(14x1.5cm)one wasdefinedat ray et2015).ray al. However,

7.30 total scale) at11°C7.30 totalscale) andconfirmedwith

N, 123°8'56.14" were collec five P olyostea robusta mbient levels ofseawaterpH shear stressesshear (F glass por ted fromted the alish Sea are approximately 7.8 experienced

Wootton andPfister2012 es. Spore release

shear flumeshear plate withclamps released andthe

2007; W), immediately transported to

(formerly Table pH aslow as7.3hasbeen

intertidal zon intertidal ig.

a decreasing gradientof

entire glass

was attached a was attached

and the time the plate and thetime plate the

that slowly that (1 1b s ). Two pH treatments

S )

was performed in , stresses and the 1

, S Pterosiphonia in stationary et al. (2000 2

and were in the

e on San Juan plate before , which ). - nd this

2.5 cm · . Water. , and )

.

This article isThis article by protected copyright. All rights reserved. exponential rise Theeffectassay. onsporedetachment ofshearstress s ineach up settlement plate, to a maximum assuming value would be some strongerthan our spores application of stress, shear weexpectedanincreasing percentage of sporesfrom todetach the in ImageJ (v. at positionand At each shearstress applied, were throughspores counted p layerve boundary shearstresses (1, increasing time attachment ofeachspecies water forstationary 35 pH reduced similar between spores the ofbothspeciestwo species; took 40 in not negatively affected by pH interaction C. and remaining using a generalized spores linearmodellogistic reg Attachmentcounted. to allowed attach connected to acamera CoolpixS3 (Nikon usingamicroscope were spores photographed (Steindorff M SXC,NewYork C. vancouveriensis

Accepted Articlevancou

b

from the attachmentfrom time the assay ;

Table S For the attachmentFor the strengthassay, attachmentFor the time low a stress assay, shear wasapplied (1 Pa) veriensis

between timeand pH;sp

seawater

1.48; U.S. National 1.48; U.S.National Institutes ofHealth, Bethesda, MD). With s each 3

- in the Supporting in the Information to

in stationary seawaterin stationary locities of0.2 -

maximum non a nd

success as a function asa success oftime wasdetermined from number the of relative to ambientc

was acute less than - 48 h P. robusta according tothe

4, 7,17and20Pa; and .

( Fig. 2

– those from

- 4

li spores

· ores that were allowed to thatwereallowed ores near regressionnear (SigmaPlot 11.0;

(Fig. 2 ms ,

a

and remainingthe photographed were and spores - and b; 1

.

, similartointertidalfield conditions ( in 300) onditions spores However, in

,

logistic logistic regression analyses of attachment C. vancouveriensis

P. a ). . and

robusta Table S multiple multiple treatments,

Reduced pHdelay

from

b . )

. Attached were spores

P. P. , 3

the magnitudethe ofthis differencewas P. ).

robusta

Although delay the in attachment robusta attach ression ression (R v. pecies wasusingpecies tested an

- for 5 52% longerto

were allowed to in set

ed there wasasignificant

see Fig.1b see for longer35 than h -

10 h R the attachment the ofboth 2

=

to maximize

0.84

3.1.2 after sporeswere then then icroscopes) attach ) - Hata 2015 , hoto ;

exposed to 0.99 represent Fig. 2 uccessive -

in analysis analysis ;

over Fig. 2

,

a were

). ing

,

This article isThis article by protected copyright. All rights reserved. biomechanical impacts see 2010 development, andreducesmacroalgal of growth ( spores composition, andabundance diversity of attachment times ofalgal strengths spores and have insensitive toreducedpH those seawater attachment to able survive highshearstresses in pH reduced conditions, the due to more rapidly (5 two distinct ofthephases attachment process. leading to anincreased frequencyobserved ofthemodal attachment strength(Fig. primarily more2d), due dislo to77% being spores 2 significantly of weakenedtheattachment strength ofdislodged regardless seawater affect attachmentdid not strength to estimatedistributions of frequency spo stress.detachment onshear Thefirst derivatives of non fitted, parameters: themaximum detached percentage ofspores c , and

Accepted Articled Padilla amd ,

that attach, despite delayed,being likelytohaveanattachment are thatis strength d Chan et al. 2015 Chan etal. , Table S Past studies Past thathave demonstrated acidified seawater delays germination, retards R

educed pHnegativelyimpacteduced f). In low pH seawater, pHIn low seawater, an f). 24%ofadditional spore the populationdislodged(Fig. –

- . perhaps increasing thelikelihoodofbeing castashoreor Gamiño al.2016 et In contrast, - 4 10 h

in the InformationSupporting ) than of spores ,

of reducedoceanpH Bradassi etal.Bradassi 2013 spore attachmentspore in

. In this manner, differing manner, effects. Inthis of on oceanacidification ,

Leal et al. 2017 Leal etal. pH

in P. P. (Table S (Table s

robusta robusta spore attachment species intwo of algae in red and macroalgal re attachmentre (Fig.2 strength ,

P. Gaitán on 4 S ; robusta ; a maximum of50

). Here, we pores of (35 ), which providedestimates oftwo Fig. 2 early early dged under moderate Pa;dged under shear(4 C. vancouveriensis - - the potentialthe toimpactfuture the 43 h

Espitia et al.2014 populations. spore ,

c was delayedgreatly inreduced pH ), C. vancouveriensis Coelho al.2000, et and

and initialdependence of the yet demonstrate negativedemonstrate adhesion, which precede

e). However, reduced pHe). However,reduced - fewer oftheses linear linear wereregressions plotted

- 55% of spores were55% ofspores washed sea outto

spores (TableS , ir ,

James 2014 al. et e

weake and

atta Hofmann al. et

pores may be f) ned ch much .

Reduced pH

2f).

s Fig. 2d),

4 and

– ;

Fig. but ,

but This article isThis article by protected copyright. All rights reserved. affected dys potentialhas the to macro orbiological chemical in nature. toclarifyneeded iftheprecise impact of acidification differences in differmucilages macroalgal widelyamong species Fletcher Callow (see and 1992) be (Quatrano and Crayton1973) possibleout the impact ofreducedoceanpHonsporephysiology viaincrea perhaps al. protonated, anionic spore mucilages (Li acidified seawater attachment affect mustmacroalgae despite ocean that changing adhere al.2017 et McCoy andKamenos 2015 andFletcher 1992 Callow colonization supersede perhaps

Accepted Article affected 1982) function, despitethe s algae

spores In summary, o ,

compromised or thedisrupting ) behighlysusceptible to pH,reduced haveuncalcified spores

. . may distributedwidely be

Because the Because the

Assuming of macroalgalof populations on spore adhesion is adhesionis on spore also expected to vary spore spore of anduncalcified both calcified macroalgae sed gel s would be

impact the cycleslife ofboth calcified and fleshy any physiological Sporeadhesion response. is adhesion (seeFig. 1; cean acidificationcomp cean

spore adhesionspore results potential viabilitypotential andresilience of mature algal thalli. chemical density density and ) interaction substrata ofmucilages (Callow with and Fletcher 1994) . , Roleda et al. 2015 , Roledaetal.

consistent with Even calcifiedmacroalgae, w

or translate tofield conditionsand substrata, natural

intercellular mucilage production

composition, curingrate composition,

viscosity (Buviscosity across speciesacross

could

to new substratato new et et al.

reduced oceanpH reduced Moorjani andJones 1972),

reduced ionic cross

cause declines in 2013), slowing potentially rates curing romised attachment spore ofred in two species ) ormay (Padilla not et al. et . conditions. Compromised sporeattachment

across species

(

2005).

Chamberlain 1976, Boney1981, hich mayhich etal.2013, (Kroeker , and

suggests on adhesion on adhesion is

seaweed diversity seaweed and - Nevertheless, wecannotrule linking and hydration of production rate T ,

hat oceanacidification hat fundamental as sulphate uptake as sulphate (Pueschel 1979) (Pueschel 1979) - Gamiño 2016 al. et that impact the the impact o species . Additionalwork is

like other like generally generally

via spore

If to the

s of spore reduced pH

on spore f ocean fitness

yielding may

(Jones (Jones

in , Leal , Leal

also also is et This article isThis article by protected copyright. All rights reserved. manuscript Carringtonand membersthe of us allowing to collect algae fromDead ManBay. troubleshooting and Herko, MollyRoberts, Michael Barsamian assistance for with data collection and Acknowledgements forgenerousdonors their contributions. c sourcing crowd funded througha Additionally, construction the ofthe settlementspore andthe shearflume apparatus was Wainwright fellowship Stephenandresearch RuthFellowship and RebeccaGuenther. to (N by Additional fundingwasprovided author the atScienceserved theFoundation. National Anyopinion, findings, are conclusions or those of 1041213) to E. Funding macroalgal populationinfluence dynamics. points and identify tipping whichdocumentedimpacts onspore adhesion beyond would producedspores bymature macroalgal thalli alongourshores. abundance

Accepted GrantSERC) toP. Discovery Article We acknowledgehelp gratefully the of O’Donnell, George, Michael Matt Michelle Funding forthi

s and donotnecessarily viewsreflect the of NationalScience the Foundation .

Carrington throughou

s project Nationalwas provided bythe Science . t this project.Wet this This materialba is F uture studies should whether address Martone Lab

T. Martone, the PhycologicalSocietyGrants in the aid ampaign (Petridish.org) to K. the Natural the Sciences and Engineering Research Council

thank the SanJuan the thank CountyLandBankfor s could compensate for increased spore mortality

sed for theirthoughtful commentsthis on

We upon work supported while E. upon worksupported also also

thank

Carolyn Friedman and Miklasz, andwetha the excessive excessive the Foundation (EF

Carrington number of nk the nk the -

of

This article isThis article by protected copyright. All rights reserved. AcceptedCeramium Chamber Third Int Chamberlain Document. Campbell bioadhesion Callow 1717. association Ugi and gelation the ofaqueous via reaction alginate. ArticleBu, H., Kj Mar coralline Bradassi Boney 518. Boney References .

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P , T. 2015 ,

, . A Ph.D R

P M ,

.

. . , K M In K , Hurd

. , G A. 2010

. . J

. J , Hepburn ,

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. This article isThis article by protected copyright. All rights reserved. ontogenetic approach.and geographic Physiological plasticity elevatedand local adaptationto pCO2 incalcareous algae:an Padilla by oceanacidification. O’Donnell naturally Paul Murray algae. W.E.1972.SporeMoorjani, attachment S.&Jones, anddevelopment insome Phycol ecological,integrating physiological, andgeochemical responsestoglobalchange. McCoy marineassemblies in gels:anatomistic simulation. Li P Lee olym ,

Accepted Article X ,

, K

. B , Leck

. Br. . - .

. & Y , ,

S Gamiño

51 S J ci . high pCO .

. P W & Mooney, D & Mooney,

. , : J Fay A.2015

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An inland sea nitrateAn inland high , , J. 2012.Alginate:properties .

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D Y . 60 . . & , Kelly 3 . :

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957 Ågren 587 ,

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966. 590. M , 9 -

, J low chlorophyll (HNLC)with region M :

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M . , This article isThis article by protected copyright. All rights reserved. Accepted Biofouling apparatus Schultz Oceanogr B.Santelices 1990Patterns of dispersaland reproduction, recruitment inseaweeds. algal andanassociatedbenthicassemblage.recruits, Effect pH acidification and ofocean fluctuations onthegrowthanddevelopment ofcoralline Roleda Article to responses thefacultative environment. Reynolds role in R Quatrano, (Rhodophyta). C Pueschel, gels. Picone

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This article isThis article by protected copyright. All rights reserved. of passed. water were creating a spores released channel over the throughworking section outflow whichthe differentsheargenerate stresses. The flumemounted ontopof thesettlement was where plate settlement plate ofmature time, thalli as weremoved across the Fig. 1: F at drivers a siteofrapidly declining pH. ocean Wootton Princeton, NJ Vogel phase:a The oceanic gel bridge in the DOM Verdugo Oceanogr macroalgal propagules:algal Cross Taylor Saccorhiza UV vshighPARphlorotannin andlow radiation on content andgermination inzoosporesof Steinhoff

Acceptedigure Captions Article , ,

S (A) Spore settlement to used access spore apparatus attachment variable amounts for D , . , ,

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J 2003 P . , Delaux F

. 55 . T dermatodea . , Alldredge S . , 640pp : . &

, Wiencke, . 66

Comparative biomechanics:Comparative life's physical world

(60 x7.6 cm)

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C . . L A. 2012 . , Wuttke, . , Azam , and (B) the shear the , and(B) flume with ,

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species inaspecies comparisons environment. turbulent , Carbon system climaticmeasurements andpotential ,

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. , This article isThis article by protected copyright. All rights reserved. Acceptedvancouveriensis Table S3: coursethe of sporeattachment assays. Table S2: treatment. isvalue assayafter after Error wascompleted. is S.E.M. for spore assay andpH each vancouveriensisCorallina Table S1: Table Captions regressions of data. detachment (F) Frequencywith shear stress. distribu Article maximum wherea x)), is the initial the detachment andbis rateofincrease ofdetachment linear regressionsofpercentdetachment spore shear of stress asfunctions (y=a*(1 attachment time =9 (n 48 h lines dashed 95% are confidence Detachment intervals. in(C) ofspores regressions (ln[Y/(1 attached)(not 1(attached) or after anapplied low stress Pa).Solid shear lines are (1 logistic Corallina strengths. Attachment times(A) sporesin of datapoints) attachmenton spore time,dislodgement, andfrequency of distribution attachment Fig. C. vancouveriensis

2: of attachment time =5 (n The effects ofco

va Generalized Generalized modelfora linear results alkalinityTotal (µ Average pH(total scal

ncouveriensis

and - Y)] = a +bX), where a is the b Y)] =a+bX),whereisthe intercept, is theslope, a Polyostea robusta. - , generated from first from derivative calculations non fitted, 11, mean11, ntrol pH(7.75ntrol

and 335 (n = mol/kgSW)measurements water fromcollected samples over - 9, mean Polyostea robusta. Polyostea robusta.

± e) andtemperature (°

S.E.M.). Linesaree - 477), eachpoint where represents one scored spore as0 tions ofspore attachment strengthsin (E) pH;

±

S.E.M.) and (D) S.E.M.) and(D)

black data Polyostea ttachment of sporesof times algal

Before up isvaluewhenassay wasset and points) andreduced pH(7.30pH; C)

robusta xponential rise

of control and low pH treatments C. vancouveriensis

(n = 614 (n = - to P. - 663) and (B) 663) and - maximum, non robusta nd Xistime, and

after 5 after P. robusta

Corallina Corallina after 35 - linear linear - - 10 h exp(b *

red red

- and

of

- of This article isThis article by protected copyright. All rights reserved. t significancethe ofparameters ineach regression, andtreatment p ofalgal spores Table S4: - Accepted Articletests comparing parameter estimates the in twotreatments andlowpH). (control

Results ofexponResults Corallina vancouveriensis ential rise ential

- to - maximum, non and Polyostea robusta. - linear regression

- Parameter p values reflect results reflect fromvalues s ondetachment of - values reflect

This article isThis article by protected copyright. All rights reserved. Accepted Article