S anda ds in Genomic Sciences (2013) 9:71-82 DOI:10.4056/sigs.4328031
The Genomic S anda ds Conso ium
Comple e genome sequence o G anulicella mallensis ype
s ain MP5ACTX8T, an acidobac e ium om und a soil
Suman R. Rawa 1, Minna K. Männis ö2, Valen in S a o oy o 3, Lynne Goodwin4, Ma
Nolan5 Lo en J. Hause 6, Mi iam Land6, Ka en Wals on Da enpo 4, Tanja Woyke5 and Max
M. Häggblom1*
1 Depa men o Biochemis y and Mic obiology, Ru ge s, The S a e Uni e si y o New
Je sey, New B unswick, New Je sey USA
2 Finnish Fo es Resea ch Ins i u e, Ro aniemi, Finland
3 Depa men o Cell Biology and Neu oscience, Ru ge s, The S a e Uni e si y o New
Je sey, Pisca away, New Je sey, USA
4 Los Alamos Na ional Labo a o y, Bioscience Di ision, Los Alamos, New Mexico, USA
5 DOE Join Genome Ins i u e, Walnu C eek, Cali o nia, USA
6 Oak Ridge Na ional Labo a o y, Oak Ridge, Tennessee, USA
*Co esponding au ho : Max M Häggblom (haggblom@sebs. u ge s.edu)
Keywo ds: cold adap ed, acidophile, und a soil, Acidobac e ia
G anulicella mallensis MP5ACTX8T is a no el species o he genus G anulicella in subdi ision
1o Acidobac e ia. G. mallensis is o ecological in e es being a membe o he dominan soil
bac e ial communi y ac i e a low empe a u es and nu ien limi ing condi ions in A c ic alpine
und a. G. mallensis is a cold-adap ed acidophile and a e sa ile he e o oph ha hyd olyzes a
sui e o suga s and complex polysaccha ides. Genome analysis e ealed me abolic e sa ili y
wi h genes in ol ed in me abolism and anspo o ca bohyd a es. These include gene mod-
ules encoding he ca bohyd a e-ac i e enzyme (CAZyme) amily in ol ed in b eakdown, u ili-
za ion and biosyn hesis o di e se s uc u al and s o age polysaccha ides including plan based
ca bon polyme s. The genome o G anulicella mallensis MP5ACTX8T consis s o a single epli-
con o 6,237,577 base pai s (bp) wi h 4,907 p o ein-coding genes and 53 RNA genes.
In oduc ion
S ain MP5ACTX8T (= ATCC BAA-1857T = DSM
23137T), is he ype s ain o he species
G anulicella mallensis [1]. The genus G anulicella,
in subdi ision 1 o Acidobac e ia, was i s de-
sc ibed by Pank a o e al. in 2010 [2].
G anulicella mallensis (mal.len' sis. N. L. em. adj.
mallensis; pe aining o i s isola ion om soil o
Malla Na u e Rese e, Kilpisjä i, Finland;
69°01’N, 20°50’E) was desc ibed along wi h o he
species o he genus G anulicella isola ed om
und a soil [1] and is one o he wo wi h se-
quenced genomes, ou o eigh alidly desc ibed
G anulicella species.
Acidobac e ia is one o he mos ubiqui ous bac e i-
al phyla ound in di e se habi a s and is abundan
in mos soil en i onmen s [3,4] including A c ic
und a soils [5,6]. Acidobac e ia a e
phylogene ically and physiologically di e se [7]
ep esen ed by 26 phylogene ic subdi isions [8] o
which only subdi isions 1, 3, 4, 8, and 10 a e de-
ined by axonomically cha ac e ized ep esen a-
i es. To da e, subdi ision 1 is comp ised o eigh
gene a: Acidobac e ium [9], Te iglobus [10,11],
Edaphobac e [12], G anulicella [1,2], Acidipila [13],
Telma obac e [14], Acidicapsa [15] and B yocella
[16]. Subdi ision 3, 4 and 10 include only one ge-
nus each, namely B yobac e [17], Blas oca ella
[18] and The mo omaculum [19], espec i ely,
while subdi ision 8 includes h ee gene a;
Holophaga [20], Geo h ix [21] and
Acan hopleu ibac e [22]. Th ee species,
‘Candida us Ko ibac e e sa ilis’ [23], ‘Candida us
Solibac e usi a us’ [23] and ‘Candida us
Chlo acidobac e ium he mophilum’ [24] ha e
been desc ibed as ‘Candida us’ axa. Acidobac e ia
a e ela i ely di icul o cul i a e wi h slow g ow h
G anulicella mallensis ype s ain MP5ACTX8T
72 S anda ds in Genomic Sciences
a es and ypically equi e up o se e al weeks o
de elop isible colonies on solid media. Ne e he-
less, he phylogene ic di e si y, ubiqui y and abun-
dance o his g oup sugges ha hey play im-
po an ecological oles in soils. The abundance o
Acidobac e ia has been ound o co ela e wi h soil
pH [25,26] and ca bon [27,28], wi h subdi ision 1
Acidobac e ia being mos abundan in sligh ly acid-
ic soils. Ou p e ious s udies ha e shown ha
Acidobac e ia domina e in he acidic und a hea hs
o no he n Finland [25,29-31]. Using selec i e iso-
la ion echniques we ha e been able o isola e se -
e al slow g owing and as idious s ains o
Acidobac e ia [1,11]. On he basis o phylogene ic,
pheno ypic and chemo axonomic da a, including
16S RNA, poB gene sequence simila i y and DNA–
DNA hyb idiza ion, s ain MP5ACTX8T was classi-
ied as a no el species o he genus G anulicella [1].
He e, we summa ize he physiological ea u es o-
ge he wi h he comple e genome sequence, anno-
a ion and da a analysis o G anulicella mallensis
MP5ACTX8T (Table 1).
Classi ica ion and ea u es
Wi hin he genus G anulicella, eigh species a e de-
sc ibed wi h alidly published names: G. mallensis
MP5ACTX8T, G. und icola MP5ACTX9T, G. a c ica
MP5ACTX2T and G. sapmiensis S6CTX5AT isola ed
om A c ic und a soil [1] and G. paludicola
OB1010T, G. pec ini o ans TPB6011T, G. osea
TPO1014T and G. agg egans TPB6028T isola ed
om sphagnum pea bogs [3]. S ain MP5ACTX8T
showed 95.5 -96.1% 16S RNA gene sequence iden-
i y o und a soil s ains, G. und icola MP5ACTX9T
(95.5%), G. sapmiensis S6CTX5AT (96.2%) and G.
a c ica MP5ACTX2T (96.1%) and 94.6 – 97.4% o G.
osea TPO1014T (94.6%), G. agg egans TPB6028T
(96.0%), G. pec ini o ans TPB6011T (96.1%), G.
paludicola OB1010T (96.5%) and G. paludicola
LCBR1 (97.4%). Phylogene ic analysis based on he
16S RNA gene o axonomically classi ied s ains
o amily Acidobac e iaceae placed G. paludicola
ype s ain OB1010 T as he closes axonomically
classi ied ela i e o G. mallensis MP5ACTX8T (Fig-
u e 1).
Figu e 1. Phylogene ic ee highligh ing he posi ion o G. mallensis MP5ACTX8T (shown in bold) ela i e o he o h-
e ype s ains wi hin SD1 Acidobac e ia. The maximum likelihood ee was in e ed om 1,361 aligned posi ions
o he 16S RNA gene sequences and de i ed based on he Tamu a-Nei model using MEGA 5 [41]. Boo s ap al-
ues >50 (exp essed as pe cen ages o 1,000 eplica es) a e shown a b anch poin s. Ba : 0.02 subs i u ions pe nu-
cleo ide posi ion. The co esponding GenBank accession numbe s a e displayed in pa en heses. S ains whose ge-
nomes ha e been sequenced, a e ma ked wi h an as e isk; G. mallensis MP5ACTX8T (CP003130), G. und icola
MP5ACTX9T (CP002480), T. saanensis SP1PR4T (CP002467), T. oseus KBS63T (CP003379) and A. capsula um ATCC
51196T (CP001472). B yobac e agg ega us MPL3 (AM162405) in SD3 Acidobac e ia was used as an ou g oup.
Rawa e al.
h p://s anda dsingenomics.o g 73
Table 1. Classi ica ion and gene al ea u es o G. mallensis s ain MP5ACTX8T acco ding o he MIGS ecommenda-
ions [32]
MIGS ID P ope y Te m E idence codea
Domain Bac e ia TAS [33]
Phylum Acidobac e ia TAS [34,35]
Class Acidobac e ia TAS [36,37]
Classi ica ion O de Acidobac e iales TAS [36,38]
Family Acidobac e iaceae TAS [34,39]
Genus G anulicella TAS [1,2]
Species G anulicella mallensis TAS [1]
Type s ain: MP5ACTX8T (= ATCC BAA-1857T = DSM
23137T)
G am s ain nega i e TAS [1]
Cell shape od TAS [1]
Mo ili y non-mo ile TAS [1]
Spo ula ion no epo ed NAS
Tempe a u e ange 4–28 °C TAS [1]
Op imum empe a u e 24–27 °C TAS [1]
pH ange 3.5–6.5 TAS [1]
Op imum pH 5 TAS [1]
Ca bon sou ce
D-glucose, mal ose, D- uc ose, D-galac ose, lac ose,
lac ulose, D-mannose, D- ibose, a inose, suc ose, e-
halose, cellobiose, D-xylose, glucu ona e
TAS [1]
MIGS-6 Habi a e es ial TAS [1]
MIGS-6.3 Salini y G ow h wi h up o 1.5% NaCl TAS [1]
MIGS-22 Oxygen equi emen ae obic TAS [1]
MIGS-15 Bio ic ela ionship ee-li ing TAS [1]
MIGS-14 Pa hogenici y non-pa hogenic NAS
MIGS-4 Geog aphic loca ion A c ic-alpine und a, Finland TAS [1]
MIGS-5 Sample collec ion 2006 TAS [1]
MIGS-4.1 La i ude 69°01’N, TAS [1]
MIGS-4.2 Longi ude 20°50’E
MIGS-4.4 Al i ude 700 m TAS [1]
aE idence codes - IDA: In e ed om Di ec Assay; TAS: T aceable Au ho S a emen (i.e., a di ec epo exis s in he
li e a u e); NAS: Non- aceable Au ho S a emen (i.e., no di ec ly obse ed o he li ing, isola ed sample, bu based
on a gene ally accep ed p ope y o he species, o anecdo al e idence). These e idence codes a e om he Gene
On ology p ojec [40].
G anulicella mallensis ype s ain MP5ACTX8T
74 S anda ds in Genomic Sciences
Mo phology and physiology
G. mallensis g ows on R2 medium (Di co) a pH
3.5–6.5 (op imum pH 5) and a +4 o +28 °C (op-
imum 24–27 °C) [1]. On R2 aga , s ain
MP5ACTX8T o ms opaque whi e mucoid colonies
wi h a diame e o app oxima ely 1 mm. Cells a e
G am-nega i e, non-mo ile, ae obic ods, app ox-
ima ely 0.5–0.7 mm wide and 0.6–1.3 mm long.
G ow h obse ed wi h up o 1.5% NaCl (w/ )
(Table 1). The cell-wall s uc u e in ul a hin sec-
ions o elec on mic og aphs o cells o
MP5ACTX8T is shown in Figu e 2.
G. mallensis u ilizes D-glucose, mal ose, cellobiose,
D- uc ose, D-galac ose, lac ose, lac ulose, D-
mannose, D- ibose, a inose, suc ose, ehalose,
D-xylose, N-ace yl-D-glucosamine, glucu ona e,
glu ama e, melezi ose and salicin, bu does no
u ilize D-a abinose, ace a e, o ma e, py u a e,
mala e, manni ol, D- o L-alanine, D-glycine, L-
leucine, L-o ni hine, gluconic acid, aspa a e,
dulci ol, bu y a e, cap oa e, ale a e, lac a e, oxa-
la e, p opiona e, uma a e, adoni ol, me hanol,
e hanol, succina e, D-so bi ol o myoinosi ol,
when g own using VL55 mine al medium wi h
100 mg yeas ex ac l-1. G. mallensis hyd olyzes
aesculin, s a ch, pec in, lamina in and lichenan,
bu no gela in, cellulose, xylan, sodium algina e,
pullulan, chi osan o chi in on R2 medium. S ains
show posi i e eac ion o acid and alkaline phos-
pha ases, leucine a ylamidase, a-chymo ypsin,
naph hol-AS-BI-phosphohyd olase, α- and β-
galac osidases, α- and β-glucosidases, N-ace yl- β-
glucosaminidase, β-glucu onidase, ypsin and
aline a ylamidase, bu nega i e o α- ucosidase,
α-mannosidase, es e ase (C4 and C8), lipase (C14)
and cys ine a ylamidase. S ain MP5ACTX8T e-
duces ni a e o ni i e. S ain MP5ACTX8T is e-
sis an o he an ibio ics e y h omycin, chlo am-
phenicol, neomycin, i ampicin, s ep omycin, gen-
amicin, polymyxin B and penicillin, bu suscep i-
ble o ampicillin, kanamycin, e acycline,
lincomycin, no obiocin and baci acin [1].
Chemo axonomy
The majo cellula a y acids in G. mallensis a e
iso-C15:0 (45.3%), C16:1ω7c (28.7%), iso-C13:0 (8.3%)
and C16:0 (8.9%). The cellula a y acid composi-
ions o s ain MP5ACTX8T we e ela i ely simila
o ha o o he G anulicella s ains wi h a y ac-
ids iso-C15:0 and C16:1ω7c being mos abundan in all
s ains. S ain MP5ACTX8T con ains MK-8 as he
majo quinone.
Genome sequencing and anno a ion
Genome p ojec his o y
G. mallensis s ain MP5ACTX8T was selec ed o
sequencing in 2009 by he DOE Join Genome In-
s i u e (JGI) communi y sequencing p og am. The
Quali y D a (QD) assembly and anno a ion we e
comple ed on Decembe 26, 2010. The comple e
genome was made a ailable on Dec. 1, 2011. The
genome p ojec is deposi ed in he Genomes On-
Line Da abase (GOLD) [42] and he comple e ge-
nome sequence o s ain MP5ACTX8T is deposi ed
in GenBank (CP003130). Table 2 p esen s he p o-
jec in o ma ion and i s associa ion wi h MIGS
e sion 2.0 [32].
Figu e 2. Elec on mic og aph o G. mallensis
MP5ACTX8T.
G ow h condi ions and genomic DNA ex ac ion
G. mallensis MP5ACTX8T was cul i a ed on R2 me-
dium as p e iously desc ibed [1]. Genomic DNA
(gDNA) o high sequencing quali y was isola ed
using a modi ied CTAB me hod and e alua ed ac-
co ding o he Quali y Con ol (QC) guidelines
p o ided by he DOE Join Genome Ins i u e [43].
Rawa e al.
h p://s anda dsingenomics.o g 75
Table 2. P ojec in o ma ion.
MIGS ID P ope y Te m
MIGS 31 Finishing quali y Finished
MIGS-28 Lib a ies used Th ee lib a ies, an Illumina GAii sho gun lib a y (GSGY), a 454 Ti anium
s anda d lib a y (GSXT, GWTA) and a pai ed end 454 (GSFP) lib a y
MIGS 29 Sequencing pla o ms 454 Ti anium s anda d, 454 Pai ed End, Illumina
MIGS 31.2 Fold co e age 18.5× (454), 213× (Illumina)
MIGS 30 Assemble s Newble , VELVET, PHRAP
MIGS 32 Gene calling me hod P odigaL, GenePRIMP
Locus Tag AciX8
Genbank ID CP003130.1
GenBank Da e o Release Decembe 1, 2011
GOLD ID Gc02349
BIOPROJECT PRJNA49957, PRJNA47903
P ojec ele ance
En i onmen al, Biogeochemical cycling o Ca bon, Bio echnological,
GEBA
Genome sequencing and assembly
The inished genome o G. mallensis MP5ACTX8T
(JGI ID 4088692) was gene a ed a he DOE Join
genome Ins i u e (JGI) using a combina ion o
Illumina [44] and 454 echnologies [45]. Fo his
genome, an Illumina GAii sho gun lib a y which
gene a ed 59,701,420 eads o aling 4537.3 Mb, a
454 Ti anium s anda d lib a y which gene a ed
136,708 eads and a pai ed end 454 lib a y wi h
an a e age inse size o 10.3 kb which gene a ed
157,336 eads o aling 172.0 Mb o 454 da a, we e
cons uc ed and sequenced. All gene al aspec s o
lib a y cons uc ion and sequencing pe o med a
he JGI can be ound a he JGI websi e [43]. The
454 Ti anium s anda d da a and he 454 pai ed
end da a we e assembled wi h Newble , e sion
2.3. Illumina sequencing da a was assembled wi h
Vel e , e sion 0.7.63 [46]. The 454 Newble con-
sensus sh eds, he Illumina Vel e consensus
sh eds and he ead pai s in he 454 pai ed end
lib a y we e in eg a ed using pa allel ph ap, e -
sion SPS - 4.24 (High Pe o mance So wa e, LLC)
[47]. The so wa e Consed [48] was used in he
inishing p ocess. The Ph ed/Ph ap/Consed so -
wa e package [49] was used o sequence assem-
bly and quali y assessmen in he subsequen in-
ishing p ocess. Illumina da a was used o co ec
po en ial base e o s and inc ease consensus
quali y using he so wa e Polishe de eloped a
JGI (Alla Lapidus, unpublished). Possible
misassemblies we e co ec ed using
gapResolu ion (Cli Han, un-published),
Dup inishe [50] o sequencing cloned b idging
PCR agmen s wi h sub-cloning. Gaps be ween
con igs we e closed by edi ing in Consed, by PCR
and by Bubble PCR (J-F Cheng, unpublished) p i-
me walks. The inal assembly is based on 74.2 Mb
o 454 da a which p o ides an a e age 18.5× co -
e age and 1318.5 Mb o Illumina da a which p o-
ides an a e age 213× co e age o he genome.

G anulicella mallensis ype s ain MP5ACTX8T
76 S anda ds in Genomic Sciences
Genome anno a ion
Genes we e iden i ied using P odigal [51] as pa
o he Oak Ridge Na ional Labo a o y genome an-
no a ion pipeline, ollowed by a ound o manual
cu a ion using he JGI GenePRIMP pipeline [52].
The p edic ed CDSs we e ansla ed and used o
sea ch he Na ional Cen e o Bio echnology In-
o ma ion (NCBI) non- edundan da abase,
UniP o , TIGRFam, P am, PRIAM, KEGG, COGs
[53,54], and In e P o. These da a sou ces we e
combined o asse a p oduc desc ip ion o each
p edic ed p o ein. Non-coding genes and miscel-
laneous ea u es we e p edic ed using RNAscan-
SE [55], RNAMMe [56], R am [57], TMHMM [58],
and signalP [59]. Addi ional gene p edic ion
analysis and unc ional anno a ion we e pe -
o med wi hin he In eg a ed Mic obial Genomes
Expe Re iew (IMG-ER) pla o m [60].
Genome p ope ies
The genome consis s o one ci cula ch omosome
o 6,211,694 bp in size wi h a GC con en o 57.8
mol% and consis s o 53 RNA genes (Figu e 3 and
Table 3). O he 4,960 p edic ed genes, 4,907 a e
p o ein-coding genes (CDSs) and 90 a e
pseudogenes. O he o al CDSs, 70.5% ep esen
COG unc ional ca ego ies and 16% consis o sig-
nal pep ides. The dis ibu ion o genes in o COG
unc ional ca ego ies is p esen ed in Figu e 3 and
Table 4.
Table 3. Genome s a is ics
A ibu e Value % o To al
Genome size (bp) 6,237,577 100
DNA coding egion(bp) 5,499,388 88.2
DNA G+C con en (bp) 3612173 57.9
DNA sca olds 1 100
To al genes 4,960 100
P o ein coding genes 4,907 98.9
RNA genes 53 1.3
Pseudo genes 90 1.8
Genes in in e nal clus e s 2,679 54
Genes wi h unc ion p edic ion 3,511 70.8
Genes assigned o COGs 3,496 70.5
Genes wi h P am domains 3,754 75.7
Genes wi h signal pep ides 797 16.1
Genes wi h ansmemb ane helices 1,291 26.0
CRISPR epea s 0 -
The o al is based on ei he he size o he genome in base pai s
o he p o ein coding genes in he anno a ed genome.
Rawa e al.
h p://s anda dsingenomics.o g 77
Figu e 3. Ci cula ep esen a ion o he ch omosome o G. mallensis MP5ACTX8T displaying ele an genome ea-
u es. F om ou side o cen e ; Genes on o wa d s and (colo by COG ca ego ies), genes on e e se s and (colo by
COG ca ego ies), RNA genes ( RNAs g een, RNAs ed, o he RNAs black), GC con en and GC skew.
G anulicella mallensis ype s ain MP5ACTX8T
78 S anda ds in Genomic Sciences
Table 4. Numbe o genes associa ed wi h gene al COG unc ional ca ego ies
Code Value %age Desc ip ion
J 167 4.32 T ansla ion, ibosomal s uc u e and biogenesis
A 2 0.05 RNA p ocessing and modi ica ion
K 332 8.58 T ansc ip ion
L 156 4.03 Replica ion, ecombina ion and epai
B 1 0.03 Ch oma in s uc u e and dynamics
D 27 0.7 Cell cycle con ol, Cell di ision, ch omosome pa i ioning
Y 0.0 0.0 Nuclea s uc u e
V 76 1.96 De ense mechanisms
T 139 3.59 Signal ansduc ion mechanisms
M 322 8.32 Cell wall/memb ane biogenesis
N 17 0.44 Cell mo ili y
Z 0.0 0.0 Cy oskele on
W 0.0 0.0 Ex acellula s uc u es
U 79 2.04 In acellula a icking and sec e ion
O 123 3.18 Pos ansla ional modi ica ion, p o ein u no e , chape ones
C 193 4.99 Ene gy p oduc ion and con e sion
G 355 9.18 Ca bohyd a e anspo and me abolism
E 258 6.67 Amino acid anspo and me abolism
F 76 1.96 Nucleo ide anspo and me abolism
H 155 4.01 Coenzyme anspo and me abolism
I 164 4.24 Lipid anspo and me abolism
P 157 4.06 Ino ganic ion anspo and me abolism
Q 125 3.23 Seconda y me aboli es biosyn hesis, anspo and ca abolism
R 527 13.62 Gene al unc ion p edic ion only
S 418 10.8 Func ion unknown
- 1,464 29.52 No in COGs
The o al is based on he o al numbe o p o ein coding genes in he genome.
Rawa e al.
h p://s anda dsingenomics.o g 79
Discussion
G anulicella mallensis ype s ain MP5ACTX8T has
he la ges genome size o 6.2 Mbp. among he
h ee und a soil s ains o subdi ision 1
Acidobac e ia [28]. Genome analysis o
G anulicella mallensis iden i ied a high abundance
o genes assigned o COG unc ional ca ego ies o
anspo and me abolism o ca bohyd a es
(9.1%) and amino acids (6.7%) and in ol ed in
cell en elope biogenesis (8.3%) and ansc ip ion
(8.6%). Fu he genome analysis e ealed an
abundance o gene modules encoding o unc-
ional ac i i ies wi hin he ca bohyd a e-ac i e
enzymes (CAZy) amily [61] in ol ed in b eak-
down, u iliza ion and biosyn hesis o ca bohy-
d a es. G. mallensis hyd olyzed complex ca bon
polyme s, including CMC, pec in, lichenin,
lamina in and s a ch, and u ilized suga s such as
cellobiose, D-mannose, D-xylose, D- ehalose. This
pa allels genome p edic ions o CDSs encoding
o enzymes such as cellulases, pec inases, algi-
na e lyases, ehalase and amylases. In addi ion,
he G. mallensis genome con ained a clus e o
genes in he neighbo hood o he cellulose syn-
hase gene (bcsAB) which included cellulase
(bscZ) (endoglucanase Y) o amily GH8, cellulose
syn hase ope on p o ein (bcsC) and a cellulose
syn hase ope on p o ein (yhjQ) in ol ed in cellu-
lose biosyn hesis. De ailed compa a i e genome
analysis o G. mallensis MP5ACTX8T wi h o he
Acidobac e ia s ains o which inished genomes
we e a ailable is epo ed in Rawa e al. [28]. The
da a hus sugges s ha G. mallensis is in ol ed in
hyd olysis, he u iliza ion o s o ed ca bohyd a es,
and in he biosyn hesis o exopolysaccha ides
om o ganic ma e and plan based polyme s in
he soil. The e o e, we in e ha s ain G. mallensis
may be cen al o ca bon cycling p ocesses in a c-
ic and bo eal soil ecosys ems.
Acknowledgemen s
The wo k conduc ed by he US Depa men o Ene gy
Join Genome Ins i u e is suppo ed by he O ice o
Science o he US Depa men o Ene gy Unde Con ac
No. DE-AC02-05CH11231. This wo k was unded in
pa by he Academy o Finland and he New Je sey
Ag icul u al Expe imen S a ion.
Re e ences
1. Männis ö MK, Rawa S, S a o oy o V, Häggblom
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