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h ps://doi.o g/10.1007/s10750-023-05181-y
REVIEW PAPER
A me a‑analysis o d ough e ec s onli e decomposi ion
ins eams
Ve ónicaFe ei a · ManuelA.S.G aça·
A u oElosegi
Recei ed: 1 Sep embe 2022 / Re ised: 1 Feb ua y 2023 / Accep ed: 16 Feb ua y 2023 / Published online: 13 Ma ch 2023
© The Au ho (s) 2023
Abs ac D ough s, o se e e educ ions o wa e
low, a e expec ed o become mo e equen and
in ense in i e s in many egions unde he ongoing
clima e change scena io. I is he e o e impo an
o unde s and s eam ecosys em unc ioning unde
d ough condi ions. We pe o med a me a-analysis o
s udies add essing d ough e ec s on li e decompo-
si ion in s eams (50 s udies con ibu ing 261 e ec
sizes) o quan i y o e all d ough e ec s on his key
ecosys em p ocess and o iden i y he main mod-
e a o s con olling hese e ec s. D ough educed
li e decomposi ion by 43% o e all, which can
impac ene gy and ma e luxes along he e o ophic
ood webs. The magni ude o d ough e ec s on li -
e decomposi ion depended on he ype o d ough
(na u al d ough > human-induced d ough ), ype
o decompose communi y (mic obes + mac oin-
e eb a es > mic obes) unde na u al d ough , cli-
ma e (wa m and humid > empe a e and Medi e -
anean) unde human-induced d ough , and on li e
iden i y. The magni ude o d ough e ec s on li e
decomposi ion also inc eased wi h he se e i y o he
d ough . The e ec s o ongoing clima e change will
likely be s onges in s eams wi h abundan sh edde s
unde going na u al d ough , especially i he s eams
become empo a y. The composi ion o he ipa ian
ege a ion may modula e he magni ude o d ough
e ec s on li e decomposi ion, which may ha e man-
agemen applica ions.
Keywo ds Ecosys em unc ioning· He e o ophic
pa hway· S eam in e mi ency· Sys ema ic e iew
In oduc ion
Li e decomposi ion is a key ecosys em p ocess in
o es s eams, whe e i sus ains aqua ic ood webs
and is pi o al in he ca bon and nu ien cycles (Wal-
lace e al., 1997; Ma ks, 2019). Once in wa e , li e
om he ipa ian ege a ion is p ocessed by mic obial
decompose s (mos ly aqua ic hyphomyce es, bu also
bac e ia) and in e eb a e sh edde s, which medi-
a e he inco po a ion o li e ca bon and nu ien s
in o seconda y p oduc ion (Hiebe & Gessne , 2002;
González & G aça, 2003).
Handling edi o : Sally A. En ekin
Supplemen a y In o ma ion The online e sion
con ains supplemen a y ma e ial a ailable a h ps:// doi.
o g/ 10. 1007/ s10750- 023- 05181-y.
V.Fe ei a(*)· M.A.S.G aça
MARE – Ma ine andEn i onmen al Sciences Cen e,
ARNET – Aqua ic Resea ch Ne wo k, Depa men o Li e
Sciences, Uni e si y o Coimb a, Calçada Ma im de
F ei as, 3000-456Coimb a, Po ugal
e-mail: [email p o ec ed]
A.Elosegi
Facul y o Science andTechnology, Uni e si y
o heBasque Coun y (UPV/EHU), PO Box644,
48080Bilbao, Spain
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Aqua ic decompose s (mic obes and in e eb a es)
and li e decomposi ion a e highly sensi i e o en i-
onmen al condi ions and li e cha ac e is ics (Boy-
e o e al., 2016; Yue e al., 2022). D ough , in pa icu-
la , is a main ac o s uc u ing s eam communi ies
and p ocesses (Rolls e al., 2012; S ubbing on e al.,
2017; Saba e e al., 2018). D ough pe iods esul in
se e e educ ion o su ace low and can e en lead o
he o al d ying in he so-called in e mi en i e s. In
ac , be ween 51 and 60% o global i e s leng h is
in e mi en (Message e al., 2021) and he o ecas ed
inc eases in ai empe a u e, wi h he consequen
inc eases in e apo anspi a ion and wa e abs ac-
ion, will exace ba e low educ ion in many a eas,
especially in a id egions (Asadieh & K akaue ,
2017). This educ ion will likely be e en s onge
in egions subjec o human-induced d ough , e.g.,
whe e wa e is wi hd awn om s eams and i e s
o i iga ion ag icul u e (Meybeck, 2003; Milliman
e al., 2008; Döll e al., 2009). In e eb a e sh ed-
de s (mos ly belonging o he o de s Plecop e a and
T ichop e a) a e especially sensi i e o he deg ada-
ion o wa e quali y (e.g., inc eases in empe a u e,
dissol ed nu ien s and conduc i i y and dec eases in
dissol ed oxygen) unde d ough condi ions (S ub-
bing on e al., 2017). Reduced low eloci y also
dec eases he ac i i ies o mic obial decompose s due
o less e icien di usion o nu ien s and oxygen a
he wa e –bio ilm in e ace (de Bee e al., 1996) and
lack o physical s imulus o he elease o spo es by
aqua ic hyphomyce es (Fe ei a & G aça, 2006; Bas-
ias e al., 2020).
Reduced low and de e io a ion o wa e quali y
unde d ough , and consequen impac s on mic o-
bial decompose s and sh edde s, o en educe li -
e decomposi ion (Saba e e al., 2018), especially
in isola ed pools and on d y s eambeds, whe e i is
ex emely slow (Langhans & Tockne , 2006; Co i
e al., 2011; Ab il e al., 2016). Li e decomposi-
ion is hus slowe in in e mi en han in pe ennial
s eams, which has been a ibu ed o legacy e ec s o
pas d y pe iods educing sh edde densi y when low
esumes (‘d ying memo y’; Da y e al., 2011). These
legacy e ec s a e likely less impo an o mic obes,
which can emain on li e e en du ing eme sed pe i-
ods al hough becoming po en ially less e icien when
wa e e u ns a e longe d y pe iods (Gonçal es
e al., 2016, 2019; A oi a e al., 2018; Mo a-Gómez
e al., 2018). The e ec s o d ough can, hus, depend
on he se e i y o low educ ion, being especially
pe asi e when he s eambed d ies ou (Langhans
& Tockne , 2006; Foulquie e al., 2015; Ab il e al.,
2016).
As sh edde s a e esponsible o s imula ing li -
e decomposi ion globally by 74% o e all (Yue
e al., 2022), li e decomposi ion media ed by he
combined ac i i ies o mic obial decompose s and
sh edde s is likely o be mo e esponsi e o d ough
han mic obial-media ed li e decomposi ion (Riedl
e al., 2013). Also, as sh edde s play a la ge ole on
he decomposi ion o li e ha is so , nu ien - ich
and has low concen a ions o ecalci an and de en-
si e compounds han on he decomposi ion o mo e
ecalci an li e (Hiebe & Gessne , 2002; Yue e al.,
2022), i is expec ed ha li e ype (e.g., lea es s.
wood) and iden i y (genus) will de e mine i s sensi-
i i y o d ough (Hill e al., 1988).
Also, s udies add essing d ough e ec s on li -
e decomposi ion use a a ie y o me hodological
app oaches and add ess a ying d ough magni udes:
s udies wi h di e en d ough se e i ies, na u al
and cul u al d ough in eal s eams, mesocosm and
expe imen al lume s udies, li e bags incuba ed in
and ou o he wa e , seasonal compa isons in sin-
gle s eams e sus compa isons be ween s eams
unde con as ing d ough egimes, be o e–a e /con-
ol–impac s udies o na u ally occu ing s. expe i-
men ally induced d ough , e c. All hese can a ec he
magni ude o d ough e ec s on li e decomposi ion
and complica e be ween-s udy compa isons (Fe ei a
e al., 2015).
We ca ied ou a me a-analysis o assess he sig-
ni icance, magni ude, and di ec ion o d ough e ec s
on li e decomposi ion in s eams. We also aimed a
de e mining he he e ogenei y among s udies and a
assessing i d ough e ec s on li e decomposi ion
depended on ype o d ough , expe imen al app oach,
se e i y o d ough , li e ype, decompose commu-
ni y, li e iden i y, and clima e. The speci ic ques-
ions and hypo heses add essed a e shown in Table1.
This me a-analysis includes 50 s udies ha con ib-
u e 261 compa isons o li e decomposi ion be ween
d ough -s essed and e e ence (non-s essed) condi-
ions. We conside ed he e ec s o all ypes o d ough
(i.e., na u al and human-induced) and add essed how
me hodological app oaches (i.e., ype o d ough ,
ype o human-induced d ough , ype o expe imen-
al d ough ), se e i y o d ough and cha ac e is ics o
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Table 1 Ques ions and hypo heses add essed, da ase s used, and loca ion o esul s
Ques ions Hypo heses Da ase used Resul
Q1: Does d ough a ec li e decomposi ion in
s eams?
H1: D ough educes li e decomposi ion due o
dec eases in in e eb a e sh edde abundance and
di e si y and educ ion o mic obial decompose
ac i i y
All Table2, Fig.3
Q2: Does he esponse o li e decomposi ion o
d ough depend on he ype o d ough (na u al s.
human-induced d ough )?
H2: Human-induced d ough has s onge e ec s on li -
e decomposi ion han na u al d ough whe e s eam
communi ies a e adap ed o low low
All Table2, Fig.3
Q3: Does he esponse o li e decomposi ion o na u al
d ough depend on he ype o compa ison (spa ial s.
empo al)?
H3: Na u al d ough has s onge e ec s when compa i-
sons a e made a spa ial scales (i.e., compa ison o
pe ennial wi h in e mi en eaches) han a empo al
scales (compa ison be ween be o e s. a e d ough
e en s) due o legacy e ec s (‘d ying memo y’) in
he la e compa ison ype (i.e., li e decomposi ion is
al eady impai ed in he be o e d ough condi ion)
Na u al d ough Table2, Fig.3
Q4: Does he esponse o li e decomposi ion o
d ough depend on he ype o human-induced
d ough (cul u al s. expe imen al d ough )?
H4: Expe imen al d ough has s onge e ec s on li e
decomposi ion han cul u al d ough due o be e
con ol o con ounding a iables in he o me han
la e condi ions
Human-induced d ough Table2, Fig.3
Q5: Does he esponse o li e decomposi ion o
d ough depend on he ype o expe imen al d ough
(simula ed wa e di e sion s. simula ed desicca ion
s. mesocosms)?
H5: Simula ed desicca ion (i.e., compa ison o
imme sed and eme sed li e samples) has s onge
e ec s on li e decomposi ion, ollowed by simula ed
wa e di e sion (i.e., dec eases on low bu gene ally
s ill we condi ions) and mesocosms (i.e., gene -
ally dec eases in low) due o di e ences in d ough
se e i y among expe imen al app oaches
Expe imen al d ough Table2, Fig.3
Q6: Does he esponse o li e decomposi ion o
d ough depend on he se e i y o d ough ?
H6: D ough e ec s on li e decomposi ion inc ease
wi h d ough se e i y since s onge educ ion in
low o highe numbe o d y days ha e mo e se e e
impac s on aqua ic communi ies (e.g., educ ion in
a ailable habi a , desicca ion) han milde educ ions
in low o ew d y days
Na u al d ough Fig.4A
Expe imen al desicca ion Fig.4B
Q7: Does he esponse o li e decomposi ion o
d ough depend on li e ype (lea es s. wood)?
H7: D ough e ec s a e s onge o he decomposi ion
o lea es whe e in e eb a es play a s onge ole han
o wood
Na u al d ough Table2, Fig.5
Q8: Does he esponse o li e decomposi ion o
d ough depend on he decompose communi y
(mic obial s. o al)?
H8: D ough e ec s a e s onge o o al li e
decomposi ion (i.e., media ed by he ac i i ies o
bo h mic obes and in e eb a es) han o mic obial-
media ed li e decomposi ion as in e eb a es will
be nega i ely a ec by bo h he d ough and he poo
mic obial condi ioning o he li e
Na u al d ough Table2, Fig.5
Human-induced d ough Table2, Fig.6
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he decomposing li e (i.e., ype, iden i y and decom-
pose communi y in ol ed) a ec ed he esponse o
li e decomposi ion o d ough (Table1). This me a-
analysis is he e o e complemen a y o a p e ious one
assessing he e ec s o human-induced d ough on
s eams (Saba e e al., 2018), including he e ec s
on li e decomposi ion (7 s udies, 41 compa isons),
bu add essing how d ough e ec s a e dependen
on egional and s eam cha ac e is ics (e.g., ain all
egime, season, s eam o de , nu ien s a us).
Me hods
Li e a u e sea ch and s udy selec ion
P ima y s udies (i.e., empi ical s udies, including
published pape s and g ay li e a u e such as Mas e o
PhD disse a ions) add essing he e ec s o d ough
on li e decomposi ion we e sea ched on May 2nd,
2022. S udies in English, published be ween Janu-
a y 1s , 1970, and Ap il 30 h, 2022 (including online
i s ), we e loca ed using Web o Science (WoS)
(da abase: Co e Collec ion; indices: Science Ci a ion
Index Expanded, Con e ence P oceedings Ci a ion
Index – Science and Book Ci a ion Index – Science).
We used he ollowing sea ch s ings (applied o he
ield ‘Topic,’ which includes i le, abs ac , keywo ds
(de ined in he s udy) and keywo ds plus (keywo ks
chosen o indexing pu poses)): (i) ‘((s eam OR
i e ) AND (d ough OR in e mi en* OR empo a y
OR epheme al) AND (decomposi ion OR p ocessing
OR b eakdown OR decay) NOT ( acies OR model*
OR mic ocosm*))’ o iden i y s udies add essing
e ec s o na u al d ough and (ii) ‘((s eam OR i e )
AND (di e sion OR abs ac ion OR sca c*) AND
(decomposi ion OR p ocessing OR b eakdown OR
decay) NOT ( acies OR model* OR mic ocosm*))’
o iden i y he s udies add essing e ec s o human-
induced d ough ( he ‘NOT’ componen aimed a
educing he numbe o non- ele an s udies). Sea ch
(i) iden i ied 2855 eco ds and sea ch (ii) iden i ied
2086 eco ds; 4751 eco ds emained a e duplica es
we e emo ed (Fig. S1).
Ti les and abs ac s we e sc eened and s udies
we e selec ed i hey add essed he e ec s o d ough
(na u al o human-induced) on ben hic decomposi ion
o li e de i ed om ee o mac ophy e species (i.e.,
lea es o wood) and incuba ed in monocul u es (i.e.,
Table 1 (con inued)
Ques ions Hypo heses Da ase used Resul
Q9: Does he esponse o li e decomposi ion o
d ough depend on li e iden i y?
H9: D ough e ec s a e s onge o he decomposi ion
o pala able li e gene a (e.g., so , nu ien - ich,
wi h low concen a ion o ecalci an and seconda y
compounds) whe e in e eb a e sh edde s play a ela-
i e la ge ole han o mo e ecalci an li e
Na u al d ough , To al decompose communi y Table2, Fig.5
Human-induced d ough Table2, Fig.6
Q10: Does he esponse o li e decomposi ion o
d ough depend on clima e?
H10: D ough e ec s a e s onge o humid clima es
whe e d ough is less common and aqua ic communi-
ies may no be adap ed o d ough s ess, han o d y
clima es whe e s eams na u ally ace se e e seasonal
educ ions in low
Na u al d ough , To al decompose communi y Table2, Fig.5
Human-induced d ough Table2, Fig.6
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no in li e mix u e) on lo ic sys ems (i.e., s eams,
i e s, ou doo a i icial channels) by compa ing a
leas one d ough -s essed and one e e ence (non-
s essed) condi ion. S udies add essing he e ec s o
d ough include hose compa ing pe ennial s. in e -
mi en ( lowing, non- lowing o d y) s eams (e.g.,
Da y e al., 2011; Ab il e al., 2016), eaches wi h
s. wi hou low o d y in in e mi en s eams (e.g.,
Co i e al., 2011; Foulquie e al., 2015; Ab il e al.,
2016), eaches wi h low in we s. d y yea s (Schlie
& Mu z, 2011), ups eam s. downs eam o dams
ha educe discha ge (Menéndez e al., 2012), wa e
di e sion (e.g., Dewson e al., 2007a, b; Dea h e al.,
2009; A oi a e al., 2017), o simula ed in e mi ency
(e.g., B ude e al., 2011; Foulquie e al., 2015).
The e o e, d ough condi ions gene ally p esen dis-
cha ge below no mal base low. S udies ha add essed
he e ec s o hyd ological changes (e.g., esul ing
om dams, seasonal looding) bu ha did no p o-
ide e idence o d ough s ess (e.g., lowe discha ge
in he a ec ed loca ion) we e no conside ed. Also,
s udies compa ing seasons, egions, o land uses ha
a e expec ed o con as in wa e a ailabili y bu ha
did no add ess d ough e ec s we e no conside ed.
A e i le and abs ac sc eening, 47 eco ds we e
kep (Fig. S1).
The ull ex was sc eened and s udies we e
selec ed o inclusion in he da abase i hey epo ed
a decomposi ion es ima e, and associa ed a iabili y
measu e (no manda o y o all s udies as his can be
impu ed i missing alues a e ew) and sample size,
o bo h d ough -s essed and e e ence condi ions;
missing in o ma ion was eques ed om au ho s
be o e a decision o impu e da a o o exclude he
s udy was made. A e accoun ing o double pub-
lica ion (i.e., when he same da a a e published in
mul iple s udies), 44 unique s udies we e included.
Addi ionally, 6 s udies known o he au ho s and ha
me he inclusion c i e ia bu we e no iden i ied in
he WoS sea ch we e added o he da abase. The inal
da abase hus included 50 s udies (Fig. S1, Tables S1
and S2).
Da a ex ac ion
S udies included in he da abase sa is ied he inclu-
sion c i e ia, bu o se e al s udies no all in o ma-
ion pe aining o li e decomposi ion did and, he e-
o e, no all da a we e ex ac ed om hese s udies
(e.g., li e decomposi ion in he hypo heic zone,
li e decomposi ion in li e mix u es, li e decom-
posi ion in he pe iod be o e d ough in be o e–a e
con ol–impac designs, li e decomposi ion a ec ed
by o he ea men s, decomposi ion o co on s ips;
TableS2). Li e decomposi ion es ima es ha com-
plied wi h inclusion c i e ia (TableS1 and S2), a i-
abili y measu es, and sample size epo ed in he ex
and in ables we e ex ac ed di ec ly, in o ma ion in
g aphs was ex ac ed wi h WebPlo Digi ize (h ps://
au om e is. io/ WebPl o Dig i ize /), and missing in o -
ma ion was eques ed om he au ho s. Fo s udies
ha epo ed li e decomposi ion o e ime (e.g.,
He bs & Reice, 1982; Boul on, 1991; Co i e al.,
2011; Schlie & Mu z., 2011), only mass emaining
o mass loss a he las sampling da e was conside ed,
and o hose ha epo ed bo h exponen ial and lin-
ea li e decomposi ion a es (e.g., Maam i e al.,
1997), he o me we e conside ed.
Va ia ion measu es we e ex ac ed as p o ided in
he p ima y s udies o by he au ho s (i.e., s anda d
de ia ion (SD), s anda d e o (SE), o 95% con i-
dence in e al (CI)). SD alues we e used di ec ly
o es ima ing he a iance associa ed wi h he e ec
size, while SE and 95% CI we e i s con e ed in o
SD. Fo s udies ha did no epo a ia ion associ-
a ed wi h li e decomposi ion (e.g., Richa dson,
1990; Boul on, 1991; Be nal, 2010; Riedl e al., 2013;
Huang e al., 2018), SD alues we e impu ed om
s udies wi h simila expe imen al designs and ha
epo ed li e decomposi ion in he same uni (Lajeu-
nesse, 2013; Appendix1).
Values ex ac ed om g aphs o impu ed may
de ia e om he eal alues, bu no conside ing hem
would ha e limi ed he analysis. Howe e , he po en-
ial bias in oduced in o he da abase by ex ac ing
da a om g aphs and by da a impu a ion was assessed
in sensi i i y analyses.
E ec size
The e ec s o d ough on li e decomposi ion we e
es ima ed as he esponse a io R, gi en by he a io
be ween he es ima e in he d ough -s essed condi-
ion (
Xd ough
) o he es ima e in he e e ence con-
di ion (
X e e ence
); analyses we e pe o med on lnR,
i.e.,
ln(
X
d ough
∕X
e e ence)
; o li e decomposi ion
exp essed as mass emaining (which a ies in he
opposi e di ec ion o mass loss o decomposi ion
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a e), he nume a o and denomina o we e swi ched
o he calcula ion o lnR (Hedges e al., 1999;
Appendix 1). R = 1 (lnR = 0) indica es no e ec o
d ough on li e decomposi ion, R < 1 (lnR < 0) indi-
ca es educ ion and R > 1 (lnR > 0) indica es s imula-
ion unde d ough . R alues can be con e ed in o
pe cen age change o ease in e p e a ion o he mag-
ni ude o he e ec (Appendix1).
The a iance associa ed wi h lnR (VlnR), needed
o weigh each e ec size in he analysis so ha mo e
p ecise e ec sizes (i.e., wi h low a iance) will be
weighed mo e and con ibu e mo e o he o e all
es ima e han less p ecise e ec sizes, was calcula ed
using he li e decomposi ion es ima e, i s SD and
sample size (Bo ens ein e al., 2009; Appendix 1).
The a iance associa ed wi h R was also used o es i-
ma e he 95% CI associa ed wi h each e ec size, so
ha R alues wi h 95% CI ha do no include 1 a e
signi ican (Appendix1).
Indi idual li e decomposi ion s udies con ibu ed
wi h mul iple e ec sizes o he da abase (2 – 36 pe
s udy) as a esul om using coa se- and ine-mesh
li e bags, se e al li e species, s eams, o d ough
ea men s. The e o e, he 50 s udies included in he
da abase con ibu ed wi h a o al o 261 e ec sizes
(Table S1 and S2). Al hough conside ing mul iple
e ec sizes pe s udy migh a ec esul s i non-inde-
pendence o e ec sizes is a p oblem, no conside ing
hem would ha e esul ed in a low numbe o e ec
sizes, which would ha e p ecluded he analysis. We
ha e, ne e heless, ca ied ou sensi i i y analyses o
assess he e ec s o non-independence o e ec sizes
on he esul s.
Mode a o a iables
Me hodological choices and en i onmen al ac-
o s may a ec he magni ude and di ec ion o he
esponse o li e decomposi ion o d ough and a e
e med ‘mode a o s’ in me a-analysis. The e o e,
in o ma ion on se e al po en ial mode a o s, acco d-
ing o ou hypo heses (Table1), was eco ded: ype o
d ough (na u al o human-induced), ype o human-
induced d ough (cul u al o expe imen al), ype o
expe imen al d ough (simula ed wa e di e sion
o desicca ion o mesocosms), ype o compa ison
being made o na u al d ough (spa ial o empo al),
pe cen age o d y days du ing he li e incuba ion
pe iod (con inuous) and pe cen age o low educ ion
(con inuous), li e ype (lea es o wood), li e
iden i y (se e al gene a), decompose communi y
in ol ed (mic obial o o al: mic obes + mac oin e e-
b a es), and clima e (se e al) (TableS3). In o ma ion
on o he a iables (e.g., wa e empe a u e, dissol ed
nu ien s, cu en eloci y, we wid h, and dep h) was
also ex ac ed, bu sample size was oo small o be
used in analyses.
S a is ical analysis
O e all e ec size
The s udies di e ed in expe imen al condi ions and,
hus, he o e all esponse o li e decomposi ion o
d ough , i.e., he g and mean e ec size, was de e -
mined using he andom-e ec s model o me a-
analysis, which conside s wo sou ces o a iance
associa ed wi h e ec sizes: wi hin-s udy a iance
(VlnR) and be ween-s udy a iance (es ima ed by he
es ic ed maximum likelihood (REML) me hod)
(Bo ens ein e al., 2009). Indi idual e ec sizes we e
weighed by he in e se o hei a iance, and he
g and mean e ec size (R) was conside ed signi ican
i i s 95% CI did no include 1. The pe cen age o
o al a iabili y ha was due o be ween-s udy a ia-
ion (I2) was also calcula ed (Bo ens ein e al., 2009).
Mode a o analyses
The e ec s o ca ego ical mode a o s on he magni-
ude and di ec ion o he esponse o li e decom-
posi ion o d ough we e assessed o subse s o he
da abase, conside ing a ailable sample size (only
mode a o le els wi h a leas h ee e ec sizes we e
es ed) and obus ness o publica ion bias. Subg oup
analysis was used o es ima e mean e ec sizes o
mode a o le els (subg oups), using he andom-
e ec s model (wi h he REML me hod o be ween-
s udy a iance) (Bo ens ein e al., 2009). He e o-
genei y was compa ed be ween (QM) and wi hin
subg oups o assess he signi icance o each mod-
e a o and subg oup. Subg oups we e signi ican i
hei 95% CI did no include 1, and wo subg oups
signi ican ly di e ed i hei 95% CI did no o e lap.
To a oid ha o he mode a o s con ound he analysis
o a gi en mode a o , ca ego ical mode a o s we e
es ed hie a chically (Fig.1).
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The e ec s o con inuous mode a o s on he
esponse o li e decomposi ion o d ough we e
assessed o subse s o he da abase by me a- eg es-
sion, using he andom-e ec s model (wi h he
REML me hod o be ween-s udy a iance) (Bo en-
s ein e al., 2009).
Sensi i i y analyses
E ec sizes we e coded as ‘p o ided’ when li -
e decomposi ion es ima es and SD we e p o ided
in nume ical o ma (i.e., shown di ec ly in he ex
o in ables o p o ided by he au ho s) o as ‘es i-
ma ed’ when alues had o be ex ac ed om g aphs
o impu ed (Table S1 and S2). The po en ial bias
in oduced in o he da abase by ex ac ing da a om
Fig. 1 Hie a chical app oach used in he subg oup analyses
showing mode a o le els wi h n ≥ 3 (mode a o le els wi h
n < 3 we e no conside ed in speci ic analyses o ha mode a-
o and a e no shown); compa ison o mode a o le els in a
subg oup analysis was done o speci ic le els o he p e ious
mode a o in he hie a chical app oach, excep i he e we e no
signi ican di e ences among le els in which case he subse-
quen analysis was made conside ing all le els o he p e ious
mode a o oge he . 1Da a om Bu ows e al. (2017) we e no
conside ed ( o al and mic obial-media ed li e decomposi-
ion da a a e shown combined); 2Da a on Cas anea, F axinus,
Ne ium, No ho agus and Ulmus li e s we e no conside ed
(n < 3); 3Da a on Ace li e was no conside ed (di e s om
mos o he li e gene a in he p e ious analysis) and da a on
cold-d y clima e was no conside ed (n < 3); 4Da a on Fagus
and Melicy us li e s we e no conside ed (n < 3); 5Da a on
Ace , Que cus and Salix li e s we e no conside ed (di e
om mos o he li e gene a in he p e ious analysis) and da a
on a id and bo eal clima es we e no conside ed (n < 3)
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g aphs and by da a impu a ion was assessed by sub-
g oup analysis compa ing he g and mean e ec
sizes o ‘p o ided’ and ‘es ima ed’ subg oups (as
desc ibed abo e o subg oup analysis). Bias would
be a conce n i he g and mean e ec size (R) would
be signi ican ly lowe (i.e., s onge e ec ) o he
‘es ima ed’ han o he ‘p o ided’ subg oup. Also,
he p e ious subg oup analyses based on he en i e
da abase we e epea ed using he ‘p o ided’ da a
only and bias would be a conce n i esul s in e p e a-
ion di e s when conside ing he en i e da abase and
when conside ing ‘p o ided’ da a only.
The po en ial e ec s o he non-independence o
e ec sizes, which esul s om each s udy con ibu -
ing wi h mul iple e ec sizes o he da abase, on he
esul s we e assessed by epea ing he analyses ( o he
ex en possible) conside ing a single e ec size pe
s udy (es ima ed in a subg oup analysis wi h ‘s udy
code’ as he mode a o and each s udy as a subg oup).
Non-independence o e ec sizes would be a p oblem
i esul s in e p e a ion based on independen e ec
sizes (i.e., one e ec size pe s udy) di e om hose
ob ained using he ull da abase (i.e., wi h mul iple
e ec s sizes pe s udy).
Publica ion bias
E idence o publica ion bias was assessed o he
en i e da abase by he unnel plo . This is a sca e
plo ha con as s he e ec sizes (lnR) wi h hei p e-
cision (SE), wi h symme ical dis ibu ion o e ec
sizes a ound he g and mean e ec size indica ing
no publica ion bias. The impac o publica ion bias
on he g and mean e ec size was assessed by he
Du al and Tweedie’s im and ill me hod (Du al &
Tweedie, 2000). This me hod es ima es a new g and
mean e ec size conside ing he ‘missing’ e ec
sizes, which a e impu ed assuming ha he unnel
plo should be symme ic. O e lap be ween he 95%
CIs o heo iginal and o he new g and mean e ec
size indica es ha he o iginal g and mean e ec size
is no s ongly a ec ed by publica ion bias.
E idence o publica ion bias in subse s o he
da abase was assessed by he Rosenbe g’s ail-sa e
numbe (N s). This alue gi es he numbe o miss-
ing e ec sizes showing an insigni ican e ec ha
would be needed o nulli y he mean e ec size, wi h
N s > 5 × n + 10 (n = numbe o e ec sizes) indica ing
ha he da ase can be conside ed obus o publica-
ion bias.
S anda d analy ic me hods we e used (g and mean
e ec size, subg oup analyses, me a- eg essions, and
publica ion bias analyses; Bo ens ein e al., 2009).
Analyses we e pe o med using OpenMEE (Wallace
e al., 2017), excep o publica ion bias analyses ha
we e pe o med using he me a o package (Viech-
baue , 2010) in RS udio (RS udio, 2012).
Resul s
Da abase
The ea lies s udy included in he da abase da es
om 1982 (He bs & Reice, 1982), and since hen,
he numbe o s udies add essing d ough e ec s on
li e decomposi ion in s eams has been accumula -
ing exponen ially eaching 50 in Ap il 2022 (Fig.
S2). The e was an a e age o 0.3 s udies/yea be o e
2000, which inc eased o 0.8 in 2000 – 2009, 2.6 in
2010 – 2019, and 3.3 s udies/yea be ween 2020 and
Ap il 2022. Mos s udies we e ca ied ou in Eu ope
(30), No h Ame ica (8) and Oceania (7) (Fig.2). Ou
o 261 compa isons o li e decomposi ion be ween
d ough -s essed and non-s essed condi ions con-
ibu ed by he selec ed s udies, 49% o igina ed om
s udies add essing na u al d ough , 43% om s udies
add essing expe imen al d ough , and 8% om s ud-
ies add essing cul u al d ough (i.e., human-induced
d ough , no caused on pu pose o he s udy) (Fig.1,
Tables S1 and S3). S udies add essing e ec s o
na u al d ough on li e decomposi ion mo e o en
pe o med spa ial (e.g., pe ennial s. in e mi en
s eam; 87%) han empo al compa isons (be o e
s. a e d ough ; 13%) (Fig. 1, Tables S1 and S3).
S udies add essing e ec s o expe imen al d ough
on li e decomposi ion mos o en used desicca ion
(imme sed s. eme sed li e bags; 64%), ollowed
by expe imen al wa e di e sion (27%) and meso-
cosm (9%) app oaches (Fig. 1, Tables S1 and S3).
Mos compa isons (91%) de i ed om lea li e and
add essed li e decomposi ion by bo h mic obes and
in e eb a es (80%) (Tables S1 and S3). Li e om
17 ee and mac ophy e gene a we e used, wi h Popu-
lus (36%), Alnus (26%), and Ph agmi es (13%) lea
li e con ibu ing mos compa isons (Tables S1).
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O e all e ec s o d ough on li e decomposi ion
The majo i y (85%) o indi idual e ec sizes lnR
we e nega i e, wi h a la ge numbe being s ongly
nega i e (Table S1), which con ibu ed o a g and
mean e ec size lnR o – 0.57 (95% CI: – 0.66 o
– 0.48) (Fig. S3). This ansla ed in o a g and mean
e ec size R o 0.57 (95% CI: 0.52 – 0.62), indi-
ca ing a signi ican educ ion o li e decomposi-
ion by 43% unde d ough condi ions (p < 0.001)
(Fig.3). The unnel plo was, howe e , asymme ic,
wi h 39 e ec sizes ‘missing’ o he le o he g and
mean e ec size (Fig. S4A), which sugges s publi-
ca ion bias. The new g and mean e ec size R es i-
ma ed by he im and ill me hod (a e impu ing
he ‘missing’ e ec sizes) was 0.49 (95% CI: 0.45
– 0.54), which sugges s a educ ion o li e decom-
posi ion unde d ough by 51%. The o iginal and he
new g and mean e ec s sizes we e, howe e , no
signi ican ly di e en ( hei 95% CIs o e lapped),
indica ing ha he g and mean e ec size based on
he da abase was no s ongly a ec ed by publica-
ions bias. In ac , he Rosenbe g’s ail-sa e num-
be was well abo e he h eshold o conside ing
he da abase obus o publica ion bias (Table 2).
The pe cen age o o al a iabili y ha is due o
be ween-s udy a ia ion was high (I2 > 99%), sug-
ges ing ha he esponse o li e decomposi ion o
d ough is a ec ed by me hodological choices and
en i onmen al ac o s.
E ec s o mode a o s on he esponse o li e
decomposi ion o d ough
The e ec o d ough on li e decomposi ion signi i-
can ly depended on he ype o d ough wi h s onge
educ ion unde na u al han unde human-induced
d ough (51% s. 35% educ ion), al hough signi i-
can in bo h cases (Table2, Fig.3). Howe e , d ough
e ec s on li e decomposi ion did no depend on he
ype o compa ison o na u al d ough (spa ial o
empo al), ype o human-induced d ough (cul u al
o expe imen al), o ype o expe imen al d ough
(wa e di e sion, desicca ion, o mesocosm s udies)
(Table 2, Fig. 3). D ough e ec s on li e decom-
posi ion depended on he se e i y o he d ough ,
wi h e ec s becoming s onge as he pe cen age
low educ ion inc eased in s udies add essing na u-
al d ough (Fig. 4A) and as he pe cen age num-
be o d y days du ing he li e incuba ion pe iod
inc eased in s udies add essing expe imen al desicca-
ion (Fig.4B). In he case o human-induced d ough ,
no signi ican ela ionship was ound be ween he
Fig. 2 Global dis ibu ion o he s udies included in he da abase (n = 50)
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decomposi ion, likely because ecological lows a e
main ained. Al hough he e is li le in o ma ion on
o he egions, we suspec ha whe e human ac i i-
ies lead o o al s eam desicca ion, i s e ec s on li -
e decomposi ion will be s ong. E ec s o d ough
a e especially s ong o o al li e decomposi ion,
sugges ing ha s eams whe e sh edde s a e abundan
will unde go a s onge educ ion o li e decomposi-
ion unde wa ming han s eams whe e li e decom-
posi ion is mos ly media ed by mic obial decom-
pose s. Also, educ ion o li e decomposi ion wi h
d ough depends on li e iden i y, which can ha e
managemen implica ions as he e ec s o d ough
may be exace ba ed o mi iga ed by changes in he
composi ion o he ipa ian ege a ion.
Acknowledgemen s We hank he many au ho s who
p o ided in o ma ion ha was no easily accessible in he
p ima y s udies and he e iewe s o hei insigh ul com-
men s. This s udy was inanced by he Po uguese Founda-
ion o Science and Technology (FCT) h ough he esea ch
p ojec STREAMECO (SFRH/BD/140761/2018), p ojec s
UIDP/04292/2020 and UIDB/04292/2020 g an ed o MARE,
and p ojec LA/P/0069/2020 g an ed o he Associa e Labo-
a o y ARNET, and by he Basque Go e nmen (IT1471-
22). Financial suppo g an ed by he FCT o VF (CEEC-
IND/02484/2018) is also acknowledged.
Au ho con ibu ions VF con ibu ed o concep ualiza ion,
p epa a ion o he li e a u e sea ch, s udy selec ion and da a
ex ac ion p o ocols, li e a u e sea ch, s udy selec ion, da a
ex ac ion, da a analysis, and w i ing. MASG con ibu ed o
concep ualiza ion, con ibu ion o he p o ocols, e ision o he
manusc ip , and app o al o i s inal e sion. AE con ibu ed o
he p o ocols, e ision o mul iple e sions o he manusc ip ,
and app o al o i s inal e sion.
Funding Open access unding p o ided by FCT|FCCN
(b-on). This s udy was inanced by he Po uguese Founda-
ion o Science and Technology (FCT) h ough he esea ch
p ojec STREAMECO (SFRH/BD/140761/2018) and he
s a egic p ojec s UIDP/04292/2020 and UIDB/04292/2020
g an ed o MARE and p ojec LA/P/0069/2020 g an ed o he
Associa e Labo a o y ARNET, and by he Basque Go e n-
men (IT1471-22). VF was inancially suppo ed by he FCT
(CEECIND/02484/2018).
Da a A ailabili y Da a used in he analyses a e p o ided in
Supplemen a y Ma e ial.
Code a ailabili y No applicable.
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in e es s.
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