Biochemis y o A c ic kelp specimens is condi ioned by he
local en i onmen
Sa ina Niedzwiedz
a,*
, Cla a Voig
a
, Sebas ian Ande sen
b
, No a Diehl
a
,
Rapha¨
elle Descˆ
o eaux
b
, Bø ge Damsgå d
b
, Kai Bischo
a
a
Ma ine Bo any, Facul y o Biology & Chemis y and MARUM, Uni e si y o B emen, 28359, B emen, Ge many
b
A c ic Biology Depa men , The Uni e si y Cen e in S alba d (UNIS), Longyea byen, NO-9171, No way
ARTICLE INFO
Keywo ds:
Pheno ypic plas ici y
Saccha ina la issima
S alba d
Tole ance
ABSTRACT
Clima e change causes empe a u e and ligh o change d as ically in A c ic jo ds, being he main d i e s o
ecosys em-enginee ing seaweeds (kelps; Lamina iales, Phaeophyceae). Clima e p ojec ions on kelps a e o en
based on s a ic pe o mance cu es, ea ing species as one homogenous uni wi h simila ole ances wi hin hei
en i e biogeog aphical ange. This migh lead o mis-ex apola ions. We assessed how A c ic kelp specimens a e
condi ioned by hei speci ic in-si u en i onmen . The e o e, we sampled Saccha ina la issima spo ophy es om
eigh jo ds along he wes coas o S alba d, No way. Analysing hei biochemical esponse a iables (pigmen
con en and composi ion; an ioxida i e ac i i y; o al ca bon and ni ogen con en ), we ound a dis inc clus-
e ing o he biochemical composi ion o S. la issima, which co ela ed signi ican ly wi h hei en i onmen .
S. la issima esponded s ongly o changes in un-o induced u bidi y, i.e., ligh a ailabili y. High ligh a ail-
abili y co ela ed wi h a signi ican educ ion o pho osyn he ic pigmen s indica ing high ligh p o ec ion.
Ne e heless, he kelps’ o al ca bon con en inc eased. The kelps’ o al ni ogen con en inc eased wi h
inc easing u bidi y, which migh be a esponse o nu ien s being washed in o he jo d by un-o . We ound no
s ess esponse o subop imal empe a u es (3 ◦C s. 7 ◦C). This is a u he indica ion o he impo ance o ligh
as a d i e o high-la i ude kelp popula ions, and he necessi y o include i in clima e p ojec ions. In conclusion,
we ound a high si e-speci ic plas ici y o A c ic S. la issima spo ophy es. This has o be conside ed when p o-
jec ing he esponses o kelps owa ds clima ic changes and local managemen ac i i ies.
1. In oduc ion
Canopy- o ming seaweeds o he o de Lamina iales, kelps, domi-
na e empe a e and A c ic ocky coas lines (S eneck e al., 2002;
We nbe g e al., 2019). Kelp o es s ha e been classi ied o be among he
mos p oduc i e ecosys ems (Pessa odona e al., 2022), ac ing as
ecosys em enginee s and ounda ion species, and hus p o iding habi a ,
nu se y g ound and ood o many associa ed o ganisms (Eckman e al.,
1989; Filbee-Dex e e al., 2019; We nbe g e al., 2019, 2024). Being
seden a y, kelps canno ac i ely escape s esso s and a e suscep ible o
en i onmen al changes. Wi hin hei gene ically se ole ance limi s,
hey ha e de eloped a ious physiological and biochemical mechanisms
o espond o he sum o en i onmen al changes, i.e., acclima isa ion
(Collie e al., 2019). The eby, hey main ain a high pe o mance, e.g.,
g ow h and ep oduc ion (pheno ypic plas ici y; King e al., 2017; Diehl
e al., 2024). Ene gy equi emen s a e lowes a a species op imum,
inc easing owa ds hei ole ance limi s, esul ing in a educed pe -
o mance (Wahl e al., 2020). A modi ica ion o he ole ance limi s o
species occu s o e gene a ions, by na u al selec ion o a ou able,
he i able ai s, i.e., adap a ion (King e al., 2017; Collie e al., 2019).
The main d i e o kelps’ la i udinal dis ibu ion is empe a u e,
while ligh has been desc ibed o de ine hei e ical dis ibu ion o e
dep h (F agkopoulou e al., 2022). Bo h empe a u e and ligh , a e
changing d as ically in A c ic jo ds wi h global clima e change (Ga uso
e al., 2020; P e idi e al., 2020, 2021; England e al., 2021; Konik e al.,
2021). V anken e al. (2021) ha e shown ha he a e o change occu s
oo as o adap i e esponses in some kelp popula ions. Gi en hei key
ecological ole, he acclima isa ion o kelps in u u e A c ic condi ions
has been subjec o many ecen s udies. Inc eased wa e empe a u es
a e p edic ed o esul in an expansion o empe a e kelp popula ion o
highe la i udes (e.g., Filbee-Dex e e al., 2019; Assis e al., 2022), while
un-o induced de e io a ion o he unde wa e ligh clima e has been
* Co esponding au ho .
E-mail add ess: [email p o ec ed] (S. Niedzwiedz).
Con en s lis s a ailable a ScienceDi ec
Ma ine En i onmen al Resea ch
jou nal homepage: www.else ie .com/loca e/ma en e
h ps://doi.o g/10.1016/j.ma en es.2025.107604
Recei ed 23 May 2025; Recei ed in e ised o m 3 Oc obe 2025; Accep ed 3 Oc obe 2025
Ma ine En i onmen al Resea ch 212 (2025) 107604
A ailable online 9 Oc obe 2025
0141-1136/© 2025 The Au ho s. Published by Else ie L d. This is an open access a icle unde he CC BY license (
h p://c ea i ecommons.o g/licenses/by/4.0/ ).
shown o esul in a shoaling o he kelp o es (e.g., Ba sch e al., 2016;
Niedzwiedz and Bischo , 2023; Düsedau e al., 2024). S udies p ojec ing
clima e change de elopmen s a e o en based on ole ances om a single
popula ion and/o single-case expe imen (Reed e al., 2011; Diehl e al.,
2024). By his app oach, i is assumed ha popula ions along he en i e
biogeog aphical ange eac as one physiological and biochemical uni
(King e al., 2017). Howe e , Benne e al. (2019) e iewed ha sus-
cep ibili ies owa ds abio ic s esso s do no only a y be ween bu also
wi hin one species, depending on a popula ion’s geog aphic loca ion
and en i onmen al his o y, which is suppo ed by se e al s udies. Fo
example, Olischl¨
age e al. (2014) ound dis inc di e ences in he
esponse o S. la issima popula ions om Helgoland and S alba d o-
wa ds he same ea men s. Liesne e al. (2020) highligh ed he
impo ance o cold seasons o he plas ici y owa ds wa ming o
Lamina ia digi a a. Gauci e al. (2022) ound ha he pe o mance o
L. digi a a spo ophy es inc eased i hei game ophy es we e
cold-p imed. Assessing he e ec s o ma ine hea wa es on no h A lan ic
kelp popula ions, Filbee-Dex e e al. (2020) concluded ha local em-
pe a u e h esholds ha e o be conside ed in p edic ions o ma ine
hea wa e consequences. Niedzwiedz e al. (2022) showed ha he sus-
cep ibili y o he kelp Saccha ina la issima owa ds expe imen al hea -
wa es s ongly depends on he ield condi ions hey expe ienced be o e
he expe imen . As cold-edge popula ions in gene al showed la ge
wi hin-species a ia ion (Benne e al., 2019), he upscaling o he e-
sponses o one, o en empe a e, popula ion o he en i e A c ic migh
pose some isk o mis-ex apola ion due o local acclima isa ion and
adap a ion p ocesses.
Saccha ina la issima is a widely dis ibu ed, cold- empe a e kelp
species, occu ing om no he n Po ugal o S alba d (A aújo e al.,
2016), wi h i s empe a u e op imum being desc ibed be ween 10 and
15 ◦C (Bol on and Lüning, 1982). Gi en i s essen ial ecological and
economic ole, i is well-s udied, se ing as model species (Diehl e al.,
2024; Sæ he e al., 2024). In his s udy, we aim o cha ac e ise he
in aspeci ic biochemical a ia ion o A c ic S. la issima, o answe he
ques ion whe he i can be ega ded as one uni . We a ge ed his
ques ion wi h an in-si u app oach in jo ds along wes e n S alba d,
No way. We mapped he abio ic d i e s o kelps ( empe a u e, ligh
a ailabili y; F agkopoulou e al., 2022) and sampled S. la issima in-
di iduals in eigh jo ds, analysing he ollowing biochemical esponses.
The kelp pigmen con en (chlo ophyll a) and composi ion ( a io o
accesso y pigmen s o chlo ophyll a (Acc:Chla); de-epoxida ion s a e o
xan hophyll cycle pigmen s (DPS; Demmig-Adams and Adams, 1996)
espond o ligh a ailabili y o mee cellula ene gy equi emen s (Blain
and Shea s, 2019). Fu he , we analysed he an ioxidan ac i i y, as
an ioxidan s we e epo ed o be a p o ec i e mechanism agains all
ypes o abio ic s esso s (Bischo and Rau enbe ge , 2012; Sha ma
e al., 2012). The o al ca bon (C) and o al ni ogen (N) con en s, as well
as hei a io (C:N) we e analysed o d aw conclusions abou he con en
o s o age compounds and nu i ional s a us o kelps a di e en sam-
pling s a ions (A kinson and Smi h, 1983). This s udy was guided by
h ee hypo heses: 1) Low-ligh in ensi ies om un-o esul in an
inc eased pigmen con en and a lowe ca bon con en . 2)
Cold- empe a u es lead o an inc eased s ess le el (high DPS and
an ioxidan ac i i y). 3) A high cellula N con en will be associa ed o
high un-o in ensi ies washing nu ien s in o he jo d
2. Ma e ial and me hods
2.1. Sampling
The A c ic A chipelago o S alba d is loca ed be ween ~77 and 80◦N
and cha ac e ised by s ong en i onmen al g adien s. The main sam-
pling campaign was conduc ed om 05 o Augus 16, 2022 by ship (MS
Spi sbe gen). S a ion 4 (Is jo den) and 5 (Eckman jo den), we e sampled
a he end o Augus (27–Augus 31, 2022). Addi ional empe a u e da a
om July and Augus 2022 a e pa o a moni o ing p og am o he
Uni e si y Cen e in S alba d (UNIS). Saccha ina la issima was collec ed
a eigh sampling s a ions along he wes coas o Spi sbe gen (Fig. 1: 1
o 8; de ailed map o di e en jo ds wi h sampling loca ions: Fig. A1).
A each sampling s a ion, simila ly sized spo ophy es (n =5) we e
collec ed om 5 ±2 m wa e dep h using a plan ake (Plan ake
19.000, acc. o Sigu d Olsen, KC Denma k, Silkebo g, Denma k).
2.2. Abio ic pa ame e s
CTD da a. One CTD p o ile pe s a ion was un in open jo d wa e
(excep o s a ion 4 and 5, whe e CTD and kelp sampling s a ions we e
he same). Wa e pa ame e s ( empe a u e [◦C], salini y [S
A
], u bidi y
[NTU]) o he wa e column we e measu ed using an RBR Maes o 3
(RBR L d., O awa, Canada). The downcas p o ile was used o moni o
an undis u bed u bidi y p o ile. The CTD was lowe ed a a speed o
app ox. 1 m s
−1
. Da a we e smoo hed by he mean o e e y me e in-
e al. Values we e epo ed as mean ±SD o he uppe 15 m o he
wa e column. Fu he CTD p o iles om c uises in 2022 be o e he
sampling pe iod we e e alua ed o demons a e p e ailing in-si u em-
pe a u e pa e ns in he espec i e jo ds.
Ligh da a. The spec ally down-welling i adiance (λ 400–700 nm)
was measu ed wi h a RAMSES-ACC-UV/VIS adiome e (T iOS Op ical
Senso , Oldenbu g, Ge many; al e na i e calib a ion) abo e he kelp
o es in wa e dep hs om he su ace un il he bo om was eached
(max 10 m; 2 o 3 p o iles pe s a ion). The i adiance o each wa e-
leng h (I
λ
) was measu ed in mW m
−2
nm
−1
and con e ed o
μ
mol
pho ons m
−2
s
−1
a e Niedzwiedz and Bischo (2023). By in eg a ing I
λ
om 400 o 700 nm, he pho osyn he ically ac i e adia ion (PAR) was
calcula ed (Niedzwiedz and Bischo , 2023). The ligh a enua ion coe -
icien (K
d
; m
−1
) was calcula ed be ween he su ace and 3 m wa e
dep h a e Hanel e al. (2001). The spec um peak (nm) a 3 m wa e
dep h was de ined as he wa eleng h wi h he maximum i adiance
ansmission h ough he wa e column.
2.3. Kelp biochemis y
A e he samples we e aken hey we e anspo ed da k, we and
cool o he labo a o y o u he p ocessing. Fo all biochemical mea-
su emen s, subsamples we e aken om each spo ophy e by cu ing
discs o pigmen analyses, 2 cm
2
in diame e , om he me is em
(app ox. basal 5 cm o he blade), wi hin 2 h o sampling. All samples
we e d ied in silica gel, which was eplaced egula ly. Samples we e
s o ed d y, cool and da k un il biochemical analyses. Biochemical
measu emen s a e based on d y weigh (DW).
Pigmen analyses. The kelp pigmen composi ion eac s o cellula
ene gy equi emen s and esponds o ligh a ailabili y (Blain and
Shea s, 2019). Pigmen concen a ions we e de e mined ollowing Koch
e al. (2015). Pe specimen, wo aliquo s o 30–100 mg powde ed ma-
e ial (n =3–5) we e ex ac ed in 1 mL 90 % ace one a 4 ◦C o 24 h in
da kness. The supe na an was il e ed and analysed by a
High-Pe o mance Liquid Ch oma og aphy (HPLC; LaCh omEli e® sys-
em, L-2200 au osample (chilled), DA-de ec o L-2450; VWR-Hi achi
In e na ional GmbH, Da ms ad , Ge many). A e W igh e al. (1991),
he pigmen s we e sepa a ed in a Sphe iso b® ODS-2 column (250 ×4.6
mm, 5
μ
m; Wa e s, Mil o d, MA, USA). Pigmen peaks we e iden i ied
and quan i ied using espec i e s anda ds. Pigmen concen a ions we e
calcula ed as
μ
g g
DW
−1
. Accesso y pigmen con en (Acc) was calcula ed as
he sum o chlo ophyll c, ucoxan hin and β-ca o ene. The a io o
accesso y pigmen s o chlo ophyll a (Acc:Chla) was calcula ed. The pool
o xan hophyll cycle pigmen s (VAZ;
μ
g g
DW
−1
) and i s a io o chlo ophyll
a was calcula ed (VAZ:Chla). The de-epoxida ion s a e o xan hophyll
cycle pigmen s (DPS; Demmig-Adams and Adams, 1996) was de e -
mined a e Colombo-Pallo a e al. (2006).
An ioxida i e ac i i y. Bischo and Rau enbe ge (2012) epo ed an-
ioxidan s o be a p o ec i e mechanism agains all ypes o abio ic
s esso s. Following he ABTS
⋅+
(2,
S. Niedzwiedz e al.
Ma ine En i onmen al Resea ch 212 (2025) 107604
2
2
′
-azino-bis-3-e hylbenz hiazoline-6-sulphonic acid, 7 mM in biDes
H2O) assay (Re e al., 1999), he kelps’ an ioxida i e ac i i y (AOA) was
de e mined. One aliquo o 50 mg powde ed ma e ial (n =3–5), was
da k-ex ac ed in 1 mL 70 % e hanol o 4 h a 47 ◦C. 10
μ
L o he su-
pe na an we e mixed wi h 1 mL ABTS
⋅+
-wo king-s anda d (abso p ion
ange: 0.740 ±0.01; 734 nm). A e an incuba ion o 6 min, he ab-
so p ion (λ 734 nm) was measu ed. AOA is epo ed as
T olox-equi alen s (TE) by calib a ing he ABTS
⋅+
-wo king-solu ion
wi h a T olox dilu ion se ies (6-hyd oxy-2,5,7,8- e ame hych o-
man-2-ca boxylic acid; 2.5 mM in 70 % / e hanol).
Ca bon and ni ogen con en . The o al ca bon (C) and o al ni ogen
(N) con en s, as well as hei a io (C:N) we e analysed o d aw con-
clusions abou he nu i ional s a us o kelp a di e en sampling s a-
ions (A kinson and Smi h, 1983). C and N con en s we e analysed using
powde ed ma e ial (2–3 mg; n =3–5). Samples we e weighed in o in
ca idges (5 ×9 mm) and combus ed a 1000 ◦C. Ace anilide was used
as s anda d (Ve a do e al., 1990). An elemen al analyse (Eu o EA 3000
Elemen al Analyse , Eu oVec o S.P.A., Milano, I aly) quan i ied he
absolu e C and N con en au oma ically. To al C and N con en s we e
exp essed in mg g
DW
−1
and as a io.
2.4. S a is ical analyses
As abio ic measu emen s we e no independen om each o he
(measu emen s o e dep h o one p o ile) and o en he limi ed numbe
o eplica es did no allow a s a is ical e alua ion, we showed he mean
and s anda d de ia ion o e dep h.
In he biochemical da a, ou lie s we e emo ed om he da ase i
hey we e classi ied as ex eme ( unc ion: iden i y_ou lie s; package:
s a ix; Kassamba a, 2023). A linea model was i on each esponse
pa ame e ( unc ion: lm; package: s a s; R Co e Team (2023) wi h
sampling s a ion as single- ixed e ec o de ec signi ican a ia ions in
kelp biochemis y. The model’s esiduals we e es ed o no mali y
(Shapi o-Wilk es , p >0.05) and homoscedas ici y (Le ene’s es , p >
0.05). As p e equisi es we e me , analysis o a iance was es ed on he
model ( unc ion: ano a; package: s a s; R Co e Team (2023). Pai wise
pe o mance ( unc ion: emmeans; package: emmeans; Len h (2024) was
used o calcula e he deg ees o eedom, wi h Tukey’s adjus men o he
p- alue. Pea son co ela ions we e analysed ( unc ion: co . es ; package:
s a s; R Co e Team (2023). A co elog am was plo ed o allow
conclusions o be d awn on he ela ionship be ween kelp biochemis y
and abio ic a ia ions ( unc ion: ggco ; package: GGally; Schloe ke
e al., 2023).
Da a we e e alua ed, plo ed and analysed in R (RS udio; V
2023.12.1 using R-4.3.2-win; R Co e Team, 2023), wi hin “ idy e se”
(Wickham e al., 2019). Maps we e c ea ed wi h ggOceanMaps
(Vih aka i, 2024).
3. Resul s
3.1. Abio ic pa ame e s
Wa e column measu emen s e ealed a dis inc g adien o all pa-
ame e s (Fig. 2A). S a ions 1 and 2 (sou h o S alba d) we e cha ac-
e ised by cold empe a u es (3.7 ±0.7 ◦C) and highes u bidi y alues
(2.8 ±2.0 NTU). A s a ions 3, 4 and 5 (mid o S alba d) empe a u es o
5.5 ±0.9 ◦C, and u bidi y alues 1.8 ±0.7 NTU we e measu ed. S a-
ions 6 and 7 in he no h o he A chipelago we e cha ac e ised by
highes empe a u es (6.6 ±0.3 ◦C) and lowes u bidi y alues (0.4 ±
0.1 NTU). Mean salini y alues ac oss all s a ions anged be ween S
A
32.0 and 33.8. Analysis o in-si u empe a u e da a om July and Augus
2022 (Fig. 2B) con i med he o e all pa e n be ween he jo ds du ing
he ime o sampling:
Mean PAR in ensi ies pe sampling s a ion a 3 m wa e dep h anged
be ween 2.3 and 137
μ
mol pho ons m
−2
s
−1
(Fig. 2C). Highes K
d
alues
we e measu ed a s a ion 2 (sou h; K
d
=1.8 ±0.4 m
-1
) and 5 (mid; K
d
=
1.0 ±0.1 m
-1
). S a ions 6, 7, and 8 (no h) clus e ed e y closely, being
cha ac e ised by lowes K
d
alues (0.2 ±0.1 m
-1
). Spec um peaks
anged be ween λ 510 and 580 nm, being highes a sampling s a ion 2 (λ
577 ±3.7 nm) and 5 (λ 582 ±1.2 nm).
Comple e CTD ( empe a u e, salini y, u bidi y) p o iles o e dep h
and he spec al esolu ion o PAR o each dep h a e p o ided as sup-
plemen a y ma e ial (Figu e A2, Figu e A3).
3.2. Kelp biochemis y
All s a is ical esul s a e summa ised in Table 1 and a e no gi en in
he ex o o e iew easons.
Pigmen s. The pigmen composi ion di e ed signi ican ly be ween
s a ions (Fig. 3A). Chlo ophyll a (
μ
g g
DW
−1
) concen a ion o samples om
Fig. 1. Sampling s a ions (1–8) o Saccha ina la issima along he wes coas o S alba d. Mean seasu ace empe a u e (SST) da a (colou g adien ) we e downloaded
om he NOAA da abase (h ps://coas wa ch.p eg.noaa.go /e ddap/, downloaded: May 08, 2024; Chambe lain, 2024), in eg a ing he SST in 2022 un il sampling
(01/01/ o Augus 16, 2022). Maps we e c ea ed wi h ggOceanMaps (Vih aka i, 2024). (Fo in e p e a ion o he e e ences o colou in his igu e legend, he eade
is e e ed o he Web e sion o his a icle.)
S. Niedzwiedz e al.
Ma ine En i onmen al Resea ch 212 (2025) 107604
3
s a ion 2 (sou h) was 50–87 % highe han in o he jo ds. Acc (
μ
g g
DW
−1
)
and VAZ (
μ
g g
DW
−1
) showed simila pa e ns (Fig. A4). The Acc:Chla
showed an inc easing end wi h highe la i ude, being signi ican ly
lowe a s a ions 1 and 2 (sou h) compa ed o s a ions 6 and 7 (no h).
While only he de-epoxida ion s a e o xan hophyll cycle pigmen s (DPS)
o s a ion 1 (sou h) was signi ican ly lowe compa ed o s a ion 6, an
o e all inc ease o DPS alues wi h highe la i ude was obse ed.
An ioxida i e ac i i y. AOA (TE mM 100 mg
DW
−1
; Fig. 3B) was signi i-
can ly a ec ed by sampling s a ions; howe e , no la i udinal end was
ound o he signi ican di e ences.
Ca bon and ni ogen con en . The C and N con en , as well as C:N we e
signi ican ly a ec ed by s a ion (Fig. 3C). Mean C con en anged
be ween 280 and 345 mg g
DW
−1
and inc eased wi h highe la i udes, wi h
he C con en o s a ion 1, 2 (sou h) being signi ican ly lowe compa ed
o s a ion 7 (no h). Mean N (7.1 and 20.1 mg g
DW
−1
) showed a dec easing
end wi h highe la i ude, al hough s a ion 7 (no h) had he o e all
highes N con en . Mean C:N (15.9 and 47.4) o s a ion 6 was signi i-
can ly highe , compa ed o he o he s a ions (excep s a ion 4).
3.3. Co ela ions
Co ela ions be ween abio ic pa ame e s and kelp esponses a e
shown in Fig. 4. O e all, a la i udinal g adien o en i onmen al pa-
ame e s was obse ed: Tempe a u e inc eased signi ican ly wi h highe
la i ude; u bidi y dec eased (no -signi ican ly) wi h highe la i ude.
Tempe a u e co ela ed nega i ely wi h u bidi y. High u bidi y alues
co ela ed wi h high K
d
alues.
Kelp pigmen concen a ions (chlo ophyll a, Acc, VAZ) co ela ed
posi i ely and pigmen a ios (Acc:Chla, DPS) co ela ed nega i ely wi h
u bidi y, he spec um peak and K
d
. DPS was ound o co ela e posi-
i ely wi h empe a u e and la i ude (signi ican ) and nega i ely wi h
u bidi y (non-signi ican ). The chlo ophyll a concen a ion, Acc and
VAZ co ela ed posi i ely wi h each o he . Al hough AOA co ela ed
posi i ely wi h PAR, his co ela ion was no signi ican .
The o al C co ela ed posi i ely wi h empe a u e and la i ude; and
nega i ely wi h u bidi y and K
d
(signi ican ). Rega ding o al N hese
co ela ions we e e e sed, hough hey we e no signi ican .
4. Discussion
The kelp Saccha ina la issima is well-s udied, being a key species in
kelp o es s along No h A lan ic coas s (A aújo e al., 2016; Diehl e al.,
Fig. 2. Abio ic pa ame e s along he S alba d la i udinal g adien . S a ions: 1 – Gnålodden, Ho nsund. 2 – Bu ge buk a, Ho nsund. 3 – Lyshamna, Bellsund. 4 –
Bjø ndalen, Is jo den. 5 – V`
o nese , Ekman jo den. 6 – Wo sleyhamna, Wood jo den. 7 – Alicehamna, Raud jo den. 8 – Y e No skøya. A) CTD da a. Tempe a u e
(◦C), Tu bidi y (NTU) o he uppe 15 m o he wa e column (mean ±SD o e dep h; n (p o iles pe s a ion) =1). No e: S a ion 8 is missing in he CTD p o iles. B)
Tempe a u e (◦C) o he uppe 15 m o he wa e column in July and Augus 2022 (mean ±SD; n (p o iles pe s a ion) =1–3). No e ha s a ions om subplo A) a e
included in he Augus mean. C) PAR: Pho osyn he ically Ac i e Radia ion (
μ
mol pho ons m
−2
s
−1
) a he su ace ( iangle; 0 m) and on 3 m dep h. K
d
(m
−1
): ligh
a enua ion coe icien be ween he wa e ’s su ace and 3 m dep h (mean ±SD; n =1–3). Spec um peak a 3 m wa e dep h: wa eleng h (nm) wi h he highes
i adiance (mean ±SD; n =1–3).
Table 1
S a is ics o biochemical pa ame e s o kelps. Resul s o Analysis o Va iance
(ANOVA) o e alua e he e ec o s a ion as single ixed e ec . Signi ican esul s
(p <0.05) a e ma ked in bold. Acc:Chla: a io o accesso y pigmen s o chlo-
ophyll a. DPS: De-epoxida ion s a e o xan hophyll cycle pigmen s. AOA:
an ioxida i e ac i i y. To al C: o al ca bon con en . To al N: o al ni ogen
con en . C:N: ca bon o ni ogen a io.
Pa ame e numDF denDF F alue p alue
Chlo ophyll a7 30 9.5 <0.001
Acc:Chla7 30 7.54 <0.001
DPS 7 27 2.9 0.02
AOA 7 26 4.8 0.001
To al C 7 31 4.5 0.001
To al N 7 28 15.9 <0.001
C:N 7 28 12.5 <0.001
No e: numDF: nume a o deg ees o eedom. denDF: denomina o deg ees o
eedom.
S. Niedzwiedz e al.
Ma ine En i onmen al Resea ch 212 (2025) 107604
4
2024), and is expec ed o expand polewa d wi h ongoing clima e change
(Filbee-Dex e e al., 2019; Assis e al., 2022). Modelling s udies a e
o en based on esponses om single popula ions o single-case expe i-
men s (Reed e al., 2011; Diehl e al., 2024). The eby, he in aspeci ic
a ia ion is being neglec ed. In his s udy, we aim o analyse he in a-
speci ic biochemical a ia ion o A c ic S. la issima. Con i ming hy-
po hesis 1, we ound ha low-ligh in ensi ies in sou he n S alba d
jo ds co ela ed wi h a highe con en o pho osyn he ic pigmen s and a
lowe ca bon con en . Con adic ing hypo hesis 2, we ound highe DPS
alues (indica ing physiological s ess) wi h wa me empe a u es. We
a ibu e his o he high-ligh a ailabili y, which inc eased along he
same g adien as ising empe a u es. Rega ding hypo hesis 3, we ound
a spa ial a ia ion in he kelps N con en , which was highe when kelps
we e in luenced by local un-o (s a ion 2; sou h) o coas al upwelling
(s a ion 7; no h).
These signi ican a ia ions in he biochemical composi ion o kelps
om di e en jo ds a e likely o esul in di e en esponses o en i-
onmen al changes as a e expec ed wi h ongoing clima e change. Hence,
ea ing kelps as one homogenous uni in clima e p ojec ions migh lead
o mis-ex apola ions, as hey migh be mo e o less ole an locally han
expec ed.
4.1. Biochemical composi ion esponds s ongly o he local en i onmen
We assessed he biochemical a ia ion o S. la issima along he wes
coas o S alba d. To be able o discuss hese a ia ions in an en i on-
men al con ex , we moni o ed empe a u e and ligh a ailabili y, as hey
ha e been desc ibed o be he mos impo an d i e s o kelp dis i-
bu ion (F agkopoulou e al., 2022). We ound spa ial di e ences o bo h
d i e s (Figs. 1 and 2).
Fjo d empe a u e pa e ns a e in luenced by cu en s (Co ie e al.,
2010). Sou he n jo ds in S alba d (especially Ho nsund; s a ion 1, 2)
a e cha ac e ised by cold empe a u es, being in luenced by he Sø kapp
Cu en ca ying A c ic wa e masses (Konik e al., 2021). Wi h highe
la i ude, we measu ed wa me empe a u es, as he in luence o he
Wes Spi sbe gen Cu en inc eased, ca ying wa m A lan ic wa e
masses (S endsen e al., 2002). All measu ed empe a u es du ing he
sampling, anged om 3 o 7◦C, which is well below he desc ibed
g ow h op imum o S. la issima o 10–15 ◦C (Bol on and Lüning, 1982).
Se e al expe imen al s udies assessed how S. la issima esponds o
empe a u es below hei op imum: Mon ei o e al. (2021) ound an
inc easing end o chlo ophyll a and a signi ican dec ease o he DPS
om 0 o 15 ◦C in young S. la issima o a empe a e popula ion (Rosco ,
B i any). Tes ing he same empe a u e ange on young spo ophy es
om a S alba d popula ion, Li e al. (2020) also ound dec easing DPS
alues wi h wa me empe a u e. As he DPS is pa o he kelps’
in e cellula s ess esponse (G oss and Jakob, 2010), hese esul s
indica e cold empe a u es in lic mo e physiological s ess han em-
pe a u es close o he op imum. We would ha e expec ed a simila
esponse in Ho nsund (s a ion 1, 2). Howe e , we ound a con a y
esponse whe e empe a u e co ela ed posi i ely wi h DPS alues. I
has o be no ed ha bo h Mon ei o e al. (2021) and Li e al. (2020)
wo ked wi h young spo ophy es om s ock cul u es, while we wo ked
on adul spo ophy es. E en hough spo ophy es o di e en ages can
show di e en esponse pa e ns o s esso s (Ma ins e al., 2017), we
do no assume ha hese di e en esponses we e due o age di e ences
o al e ed he mal ole ances o A c ic popula ions. In e p e ing he
biochemical composi ion o popula ions in his s udy, i has o be
conside ed ha wa me empe a u es co ela ed wi h less un-o and a
clea e wa e column (i.e., lowe u bidi y, K
d
; Figs. 2 and 4). Hence, we
assume ha high DPS alues did no p edominan ly espond o changes
in empe a u e bu a he coun e ac ed high-ligh s ess (no e ha he
Fig. 3. Kelp biochemis y along he S alba d la i udinal g adien (mean ±SD; n =3–5). S a ions: 1 – Gnålodden, Ho nsund. 2 – Bu ge buk a, Ho nsund. 3 –
Lyshamna, Bellsund. 4 – Bjø ndalen, Is jo den. 5 – V`
o nese , Ekman jo den. 6 – Wo sleyhamna, Wood jo den. 7 – Alicehamna, Raud jo den. 8 – Y e No skøya.
Di e en le e s indica e signi ican (p <0.05) di e ences be ween s a ions. Some e o ba s a e wi hin he diame e o he symbol. A) Chl a: chlo ophyll a con-
cen a ion (
μ
g g
DW
−1
). Acc:Chla: a io o accesso y pigmen s o chlo ophyll a. DPS: de-epoxida ion s a e o xan hophyll cycle pigmen s. В) AOA: An ioxida i e ac i i y
(TE mM 100 mg
DW
−1
). C) C: o al ca bon con en (mg g
DW
−1
). N: o al ni ogen con en (mg g
DW
−1
). C:N: ca bon o ni ogen a io.
S. Niedzwiedz e al.
Ma ine En i onmen al Resea ch 212 (2025) 107604
5
co ela ion is no signi ican ; Fig. 4). High ligh a ailabili y migh
sa u a e he pho osyn he ic elec on anspo chain and deple e
educ i e equi alen s (Bischo and Rau enbe ge , 2012). Excessi e
elec ons con ibu e o o m eac i e oxygen species (ROS), which can
des oy c i ical mac omolecules (Sha ma e al., 2012). DPS unc ions as
p o ec i e mechanism o dissipa e excessi e ene gy (Demmig-Adams
and Adams, 1996), he eby educing he po en ial o ROS o ma ion.
Kelps being exposed o highe ligh a ailabili y we e u he cha ac-
e ised by a signi ican ly lowe chlo ophyll a con en , mi iga ing
high-ligh s ess as less elec ons en e he elec on anspo chain (Ki k,
2011). The a io o Acc:Chla u he con i med his esponse pa e n.
Acc:Chla co ela ed nega i ely wi h he spec um peak, indica ing ha
he pigmen composi ion did no espond o un-o -induced spec um
shi s (Fig. 2, A3) in o de o close he g een gap o chlo ophyll a (S omp
e al., 2007). As we ound Acc:Chla o inc ease wi h highe ligh a ail-
abili y (Figs. 3 and 4), he educ ion o he ligh ha es ing an enna
complex a pho osyn he ic eac ion cen es u he p e en ed elec ons
being ansmi ed in o he elec on anspo chain (Falkowski and
Ra en, 2007). I has o be no ed ha we measu ed a maximum absolu e
PAR alue o ~140
μ
mol pho on m
−2
s
−1
a 3 m wa e dep h (Fig. 2C),
which is well below PAR alues ha kelps expe ience in o he egions, e.
g., in he Ge man Bigh (400–500
μ
mol pho ons m
−2
s
−1
, S ahl e al.,
2024). I also has o be conside ed ha poin measu emen s o absolu e
PAR alues a e subjec o ansien wea he condi ions (pe s. obs.), such
as clea sky (s a ion 4) s. cloud co e (s a ions 6, 7, 8). Niedzwiedz e al.
(2024) a gued ha ligh s ess a ela i ely low ligh in ensi ies is caused
by long pho ope iod and he lack o a eco e y phase. Ou in-si u s udy
suppo s hei expe imen al indings.
This line o a gumen also u he adds o he indings o Diehl e al.
(2021). They desc ibed o e all low DPS alues and excep ionally high
g ow h a es as a esponse o expe imen al empe a u es be ween 0 and
6 ◦C (including in-si u condi ions). To be able o compa e expe imen s
ac oss la i udes, hey exposed he S alba d samples o a 16:8 h L:D cycle.
We hypo hesise ha he o e all, empe a u e-independen inc ease in
pe o mance Diehl e al. (2021) ound in kelps om he A c ic is due o a
mo e op imal ligh egime and a eco e y om high-ligh s ess. These
indings a e a u he indica ion o he impo ance o ligh , shaping he
pe o mance and la i udinal dis ibu ion o high-la i ude kelps (Diehl
e al., 2024a).
We did no de ec he same s ess esponse in AOA as o DPS. We
sugges ha high-ligh s ess was al eady e ec i ely mi iga ed by
changes in he pigmen composi ion o he pho osys em, p e en ing
o e all highe cellula s ess esponse. As he AOA was signi ican ly
in luenced by he sampling s a ions (Table 1), we assume ha pa ame-
e s no quan i ied in his s udy migh ha e igge ed i .
We ound a clea end o inc easing o al C con en wi h highe
la i udes, which is co ela ing signi ican ly wi h wa me empe a u es
and highe ligh a ailabili y (lowe u bidi y; lowe kd). A highe C
con en migh indica e mo e s o age compounds, e.g., manni ol and
lamina in (G ai e al., 2016). Scheschonk e al. (2019) showed ha
S. la issima’s s o age compounds we e deple ed by 96 % a e h ee
mon hs o Pola Nigh . A educed ca bon con en a he end o he A c ic
summe pe iod migh inc ease he likelihood o S. la issima ha ing an
o e all nega i e ca bon balance, i.e., migh un in o s a a ion du ing
win e mon hs. To be able o p edic he ac ual kelp loss due o
un-o -induced low-ligh in ensi ies, he cumula i e annual PAR a ail-
abili y would ha e o be moni o ed along jo d g adien s. Cas o de la
Gua dia e al. (2023) ound ha kelps need a minimum o 49 mol
pho ons m
−2
y
−1
o o m a s able popula ion.
C:N a ios abo e 20 indica ed ha kelps om all sampling s a ions
we e N limi ed (Fig. 3; A kinson and Smi h, 1983). In summe , A c ic
wa e masses a e s a i ied by s ong empe a u e and salini y g adien s
(Fig. A2), leading o he absence o e ical mixing and e-supply o
deep-wa e nu ien s (Co ie e al., 2010). The o al N con en showed a
dec easing end wi h highe la i ude (Fig. 3). We a ibu e his o he
highe un-o in luence in sou he n S alba d jo ds (highe u bidi y;
Fig. 2), ca ying nu ien s om e es ial sou ces (McGo e n e al.,
2020). Kelps om s a ion 7 a e an excep ion, being cha ac e ised by a
high N con en . A s a ion 7, empe a u e g adien s o e dep h we e e y
weak (Fig. A2), showing almos no a ia ion wi hin he uppe 15 m o
he wa e column (Fig. 2, A2). As salini y and u bidi y di e ences we e
likewise no e y p onounced, we sugges ha s a ion 7 migh ha e been
in luenced by local upwelling, leading o a local esupply o nu ien s. As
C:N a ios e lec ood quali y o kelps (Lowman e al., 2022), his di -
e ence migh change he e iciency o ene gy ans e o highe ophic
le els (Lowman e al., 2022) and he ecosys em’s nu ien cycling.
4.2. Saccha ina la issima specimens a e no one homogenous uni
Bol on and Lüning (1982) desc ibed he empe a u e op imum o
S. la issima be ween 10 and 15 ◦C. Following he esul ing pe o mance
cu e, his indica es an inc easing s ess le el as empe a u es de ia e
om he op imum. In e ac ing wi h high ligh , we ound he opposi e
end; s ess le els inc eased wi h empe a u es close o he species
op imum. P obably empe a u es close o he op imum migh al eady
ha e mi iga ed high-ligh s ess, as was expe imen ally shown by
Niedzwiedz e al. (2024) and Diehl e al. (2024a). Fu he , we ound
kelps in mos jo ds o be ni ogen limi ed. Nu ien limi a ion was
shown o weaken kelps, making hem mo e suscep ible o o he
s esso s, such as UV adia ion (Da ison e al., 2007). Ou da a sugges
ha he na u al a ia ion in p esen -day A c ic jo ds leads o signi ican
a ia ions in he biochemical composi ion o kelps and consequen ly
al e ed suscep ibili ies owa ds s esso s.
Compa ing he biochemical composi ion o S. la issima ac oss
Eu ope, Diehl e al. (2023) ound nei he a clea la i udinal g adien no
a clus e ing o popula ions d i en by abio ic condi ions. In hei s udy,
Fig. 4. Linea dependency be ween kelp esponse pa ame e s and abio ic
condi ions. Colou scale: Pea son co ela ion co-e icien ( ). PAR: Pho osyn-
he ically Ac i e Radia ion. K
d
: ligh a enua ion coe icien . Acc: accesso y
pigmen s. VAZ: pool o xan hophyll cycle pigmen s. Acc:Chla: a io o accesso y
pigmen s o chlo ophyll a. DPS: de-epoxida ion s a e o xan hophyll cycle
pigmen s. AOA: an ioxida i e ac i i y. C: o al ca bon con en . N: o al ni ogen
con en . C:N: ca bon o ni ogen a io. As e isks: signi icance o co ela ion (*:
p <0.05; **: p <0.01; ***: p <0.001). (Fo in e p e a ion o he e e ences o
colou in his igu e legend, he eade is e e ed o he Web e sion o
his a icle.)
S. Niedzwiedz e al.
Ma ine En i onmen al Resea ch 212 (2025) 107604
6
hey ocussed on empe a u e and salini y as main d i e s, no
measu ing ligh a ailabili y o ligh -dependen esponses (e.g., pig-
men s). Based on he esul s o ou s udy, we hypo hesise ha he in-
clusion o ligh a ailabili y migh ha e esul ed in a mo e dis inc
clus e ing o sampling s a ions; especially he obse ed phlo o annin
concen a ions in he s udy o Diehl e al. (2023) indica ed he signi i-
cance o ligh on he biochemis y o kelps and popula ion di e ences.
4.3. Me hodological conside a ions
Ou s udy was ca ied ou in Augus 2022 and consis s o empo al
and spa ial poin measu emen s o bo h abio ic and bio ic samples.
Hence, he alues epo ed he e do no e lec he seasonal de elopmen
o condi ions o pa ame e s and a e also no ep esen a i e o he en i e
jo ds, which a e known o be cha ac e ised by s ong en i onmen al
g adien s (Mei e e al., 2017; Sej e al., 2024). Howe e , sa elli e da a o
he jo d bounda y condi ions (Fig. 1) and addi ional empe a u e
measu emen s o p e ious c uises (Fig. 2B), con i m he desc ibed
empe a u e pa e ns du ing he kelp sampling. Addi ional da a on he
ligh a ailabili y we e no a ailable o us.
We mainly ocussed on empe a u e and ligh a ailabili y, as hey
we e desc ibed o ac as main d i e s o he dis ibu ion o kelps
(F agkopoulou e al., 2022). Consis en wi h ha , we ound se e al
signi ican co ela ions o he biochemical pa ame e s wi h empe a u e
and ligh a ailabili y (Fig. 4). In expe imen al s udies in he labo a o y,
biochemical esponses o en i onmen al condi ions in kelps we e
de ec ed wi hin a ew days (e.g., Diehl e al., 2021; Niedzwiedz e al.,
2024), which is why we conside i plausible ha he biochemis y
esponded o en i onmen al pa ame e s. Long- e m moni o ing p o-
g ams o abio ic condi ions a e necessa y o be able o d aw conclusions
on empo al a ia ions.
While he absolu e epo ed alues in his s udy a e a snapsho o he
si ua ion du ing he sampling, we showed ha he in aspeci ic a i-
abili y o S. la issima along he wes coas o S alba d is high and ha he
species canno be ega ded as one homogeneous uni .
4.4. Concluding ema ks
In his moni o ing s udy, we compa ed S. la issima om eigh A c ic
jo ds. Based on hei a ia ion in biochemical signa u es hey canno be
ega ded as one homogenous uni in clima e p ojec ions. We ela ed he
biochemical esponses o di e ences in empe a u e and ligh a ail-
abili y o hei local en i onmen .
The eby, ou s udy highligh s he impo ance o conside ing ligh as
a d i e o he biogeog aphical dis ibu ion o kelps a hei cold-
dis ibu ion edge. While he high biochemical a ia ion we ound in-
dica es a high pheno ypic plas ici y o S. la issima, he signi ican co -
ela ions wi h abio ic en i onmen al ac o s also sugges s ha he kelps
a e condi ioned by hei local en i onmen s (Fig. 4). Hence, esponses
om single loca ion ha e o be ex apola ed wi h ca e and local en i-
onmen al condi ions ha e o be conside ed.
CRediT au ho ship con ibu ion s a emen
Sa ina Niedzwiedz: W i ing – e iew & edi ing, W i ing – o iginal
d a , Visualiza ion, Valida ion, Me hodology, In es iga ion, Fo mal
analysis, Da a cu a ion, Concep ualiza ion. Cla a Voig : W i ing – e-
iew & edi ing, Valida ion, In es iga ion, Fo mal analysis, Da a cu a-
ion. Sebas ian Ande sen: W i ing – e iew & edi ing, Valida ion,
Me hodology, In es iga ion. No a Diehl: W i ing – e iew & edi ing,
Valida ion, Concep ualiza ion. Rapha¨
elle Descˆ
o eaux: W i ing – e-
iew & edi ing, Valida ion, So wa e, In es iga ion. Bø ge Damsgå d:
W i ing – e iew & edi ing, Valida ion, Supe ision, Resou ces, Funding
acquisi ion. Kai Bischo : W i ing – e iew & edi ing, Valida ion, Re-
sou ces, Funding acquisi ion, Concep ualiza ion.
Financial Suppo
The ield wo k on S alba d was g an ed unde RiS numbe : 10890.
This s udy was conduc ed in he ame o he p ojec FACE-IT (The
Fu u e o A c ic Coas al Ecosys ems – Iden i ying T ansi ions in Fjo d
Sys ems and Adjacen Coas al A eas) and SEA-Ques e (Blue Ca bon
P oduc ion, Expo and Seques a ion in Eme ging Pola Ecosys ems).
FACE-IT has ecei ed unding om he Eu opean Union’s Ho izon 2020
esea ch and inno a ion p og amme unde g an ag eemen No.
869154. SEA-Ques e has ecei ed unding om he Eu opean Union’s
Ho izon Eu ope esea ch and inno a ion p og amme unde G an
Ag eemen No. 101136480. Views and opinions exp essed a e howe e
hose o he au ho s only and do no necessa ily e lec hose o he
Eu opean Union. Nei he he Eu opean Union no he g an au ho i y
can be held esponsible o hem.
Decla a ion o compe ing in e es
The au ho s decla e no con lic o in e es .
Acknowledgemen s
The au ho s hank he Uni e si y Cen e in S alba d (UNIS) and c ew
om MS Spi sbe gen (HX Expedi ions) o assis ance du ing ield wo k
and o logis ical, labo a o y and adminis a i e suppo . We hank
B i a Iken and And eas Suchopa o hei suppo du ing biochemical
analyses.
Appendix A. Supplemen a y da a
Supplemen a y da a o his a icle can be ound online a h ps://doi.
o g/10.1016/j.ma en es.2025.107604.
Da a a ailabili y
All p ima y da a suppo ing his s udy a e openly a ailable:
h ps://doi.pangaea.de/10.1594/PANGAEA.968466
h ps://doi.pangaea.de/10.1594/PANGAEA.968464
h ps://doi.pangaea.de/10.1594/PANGAEA.968642.
As mean empe a u e da a o July/Augus 2022 we e included as
me a-analysis in his s udy and a e pa o ano he s udy, hey will be
published along wi h i . Meanwhile con ac Rapha¨
elle Descˆ
o eaux
([email p o ec ed]) o mo e in o ma ion.
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