S anda ds in Genomic Sciences (2014) 9:449-461 DOI:10.4056/sigs.4648353
The Genomic S anda ds Conso ium
Comple e genome sequence o G anulicella und icola
ype s ain MP5ACTX9T, an Acidobac e ia 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 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 espondence: Max M Häggblom (haggblom@sebs. u ge s.edu)
Keywo ds: cold adap ed, acidophile, und a soil, Acidobac e ia
G anulicella und icola s ain MP5ACTX9T is a no el species o he genus G anulicella in
subdi ision 1 Acidobac e ia. G. und icola is a p edominan membe o soil bac e ial
communi ies, ac i e a low empe a u es and nu ien limi ing condi ions in A c ic alpine
und a. The o ganism is a cold-adap ed acidophile and a e sa ile he e o oph ha hyd o-
lyzes 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, including
gene modules encoding o he ca bohyd a e-ac i e enzyme (CAZy) amilies o he b eak-
down, u iliza ion and biosyn hesis o di e se s uc u al and s o age polysaccha ides such as
plan based ca bon polyme s. The genome o G. und icola s ain MP5ACTX9T consis s o
4,309,151 bp o a ci cula ch omosome and i e mega plasmids wi h a o al genome con-
en o 5,503,984 bp. The genome comp ises 4,705 p o ein-coding genes and 52 RNA
genes.
In oduc ion
The s ain MP5ACTX9T (=ATCC BAA-1859T =DSM
23138T) is he ype s ain o G anulicella
und icola [ un.d i.co’la. N.L. n. und a, und a, a
cold eeless egion; L. masc. su ix -cola ( om L. n.
incola) dwelle ; N.L. n. und icola und a dwelle ]
ha was isola ed om soil a he Malla Na u e Re-
se e, Kilpisjä i, Finland; 69°01’N, 20°50’E) and
desc ibed along wi h o he species o he genus
G anulicella isola ed om und a soil [1].
Acidobac e ia is a phylogene ically and physiolog-
ically di e se phylum [2,3], he membe s o which
a e ubiqui ously ound in di e se habi a s and a e
abundan in mos soil en i onmen s [4,5] includ-
ing A c ic und a soils [6,7]. Acidobac e ia a e el-
a i ely di icul o cul i a e, as hey ha e slow
g ow h a es. To da e only subdi isions 1, 3, 4, 8,
10 and 23 Acidobac e ia a e de ined by axonomi-
cally cha ac e ized ep esen a i es [8-23] as well
as h ee ‘Candida us’ axa [24,25]. The phyloge-
ne ic di e si y, ubiqui y and abundance o his
g oup sugges ha hey play impo an ecological
oles in soils. The abundance o Acidobac e ia co -
ela es wi h soil pH [26,27] and ca bon [28,29],
wi h subdi ision 1 Acidobac e ia being mos
abundan in sligh ly acidic soils. Acidobac e ia,
including membe s o he gene a G anulicella and
Te iglobus, domina e he acidic und a hea hs o
no he n Finland [26,30-32]. Using selec i e
G anulicella und icola ype s ain MP5ACTX9T
450 S anda ds in Genomic Sciences
isola 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 MP5ACTX9T was
classi ied as a no el species o he genus
G anulicella [1]. He e, we summa ize he physio-
logical ea u es oge he wi h he comple e ge-
nome sequence, anno a ion and da a analysis o
G anulicella und icola s ain MP5ACTX9T.
Classi ica ion and ea u es
Wi hin he genus G anulicella, eigh species a e
desc ibed wi h alidly published names: G.
mallensis MP5ACTX8T,G. und icola MP5ACTX9T, G.
a c ica MP5ACTX2T,G. sapmiensis S6CTX5AT iso-
la ed om A c ic und a soil [1] and G. paludicola
OB1010T, G. paludicola LCBR1, G. pec ini o ans
TPB6011T ,G. osea TPO1014T ,G. agg egans
TPB6028T isola ed om sphagnum pea bogs [2].
S ain MP5ACTX9T sha es 95.5 - 97.2% 16S RNA
gene iden i y wi h und a soil s ains G. mallensis
MP5ACTX8T (95.5%), G. a c ica MP5ACTX2T
(96.9%), G. sapmiensis S6CTX5AT (97.2%) and
95.2 – 97.7% iden i y wi h he sphagnum bog
s ains, G. pec ini o ans TPB6011T (97.7%), G.
osea TPO1014T (97.2%), %), G. agg egans
TPB6028T (96.8%), G. paludicola LCBR1 (95.9%),
and G. paludicola s ain OB1010T (95.3%), which
we e isola ed om sphagnum pea . Phylogene ic
analysis based on he 16S RNA gene o axonomi-
cally classi ied s ains o amily Acidobac e iaceae
placed G. osea ype s ain T4T (AM887759) as he
closes axonomically classi ied ela i e o G.
und icola s ain MP5ACTX9T (Table 1, Figu e 1).
Figu e 1. Phylogene ic ee highligh ing he posi ion o G. und icola MP5ACTX9T (shown in bold) ela i e o he
o he ype s ains wi hin subdi ision1 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 [42]. Boo -
s ap alues >50 (exp essed as pe cen ages o 1,000 eplica es) a e shown a b anch poin s. Ba : 0.01 subs i u ions
pe nucleo ide posi ion. The co esponding GenBank accession numbe s a e displayed in pa en heses. S ains
whose genomes 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 451
Table 1. Classi ica ion and gene al ea u es o G. und ic ola s ain MP5ACTX9T
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 [37,38 ]
Family Acidobac e iaceae TAS [35,39 ]
Genus G anulicella
TAS [1,40]
Species G anulicella und icola TAS [1]
Type s ain: MP5ACTX9T
(ATCC BAA-1859T = DSM 2 3138T)
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 21–24 °C TAS [1]
pH ange; Op imum 3.5–6.5; 5 TAS [1]
Ca bon sou ce
D-glucose, mal ose, cellobiose, D- uc ose, D-galac ose,
lac ose, lac ulose, D-mannose, suc ose, ehalose, D-xylose,
a inose, N-ace yl-D-glucosamine, glu ama e
TAS [1]
MIGS-6 Habi a e es ial, und a soil TAS [1]
MIGS-6.3 Salini y No g ow h wi h >1.0% NaCl (w/ ) 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 hogen NAS
MIGS-4 Geog aphic loca ion Malla Na u e Rese e, 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 TAS [1]
MIGS-4.4 Al i ude 700 m TAS [1]
a E 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 [41].
G anulicella und icola ype s ain MP5ACTX9T
452 S anda ds in Genomic Sciences
Mo phology and physiology
G. und icola cells a e G am-nega i e, non-mo ile,
ae obic ods, app oxima ely 0.5 μm wide and 0.5 –
1.8 μm long. Colonies on R2A aga a e pink, ci cu-
la , con ex and smoo h. G ow h occu s a +4 o
28°C and a pH 3.5-6.5 wi h an op imum a 21-24°C
and pH 5 (Fig. 2). Geno ypic analyses, including low
poB gene sequence simila i y and pheno ypic
cha ac e is ics clea ly dis inguished s ain
MP5ACTX9T om o he G anulicella spe-
cies/s ains, leading us o conclude ha
MP5ACTX9T ep esen s a no el species o he genus
G anulicella, o which he name G anulicella
und icola sp. no . was p oposed [1].
S ain MP5ACTX9T hyd olyzed complex o simple
ca bon subs a es [1] which include complex poly-
saccha ides like aesculin, pec in, lamina in, s a ch
and pullulan, bu no gela in, cellulose, lichenan,
sodium algina e, xylan, chi osan o chi in. S ain
MP5ACTX9T also u ilized he ollowing suga s as
g ow h subs a es: D-glucose, mal ose, cellobiose,
D- uc ose, D-galac ose, lac ose, lac ulose, D-
mannose, suc ose, ehalose, D-xylose, a inose, N-
ace yl-D-glucosamine, glu ama e and gluconic acid.
Enzyme ac i i ies epo ed o he s ain
MP5ACTX9T include acid phospha ase, es e ase (C4
and C8), leucine a ylamidase, aline a ylamidase,
α-chymo ypsin, ypsin, naph hol-AS-BI-
phosphohyd olase, α- and β-galac osidases, α- and
β-glucosidases, N-ace yl- β-glucosaminidase, β-
glucu onidase, α- ucosidase and α-mannosidase
bu nega i e o alkaline phospha ase and lipase
(C14). S ain MP5ACTX9T is esis an o ampicillin,
e y h omycin, chlo amphenicol, neomycin, s ep-
omycin, e acycline, gen amicin, baci acin,
polymyxin B and penicillin, bu suscep ible o i-
ampicin, kanamycin, lincomycin and no obiocin.
Chemo axonomy
The majo cellula a y acids in G. und icola a e
iso-C15:0 (46.4%), C16:1ω7c (35.0%) and C16:0 (6.6%).
The cellula a y acid composi ion o s ain
MP5ACTX9T was simila o ha o o he
G anulicella s ains wi h a y acids iso-C15:0 and
C16:1ω7c being mos abundan in all s ains. S ain
MP5ACTX9T con ains MK-8 as he majo quinone
and also con ains 4% o MK-7.
Genome sequencing and anno a ion
Genome p ojec his o y
G. und icola s ain MP5ACTX9T was selec ed o
sequencing in 2009 by he DOE Join Genome Ins 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 May 24, 2010. The GenBank Da e o
Release was Feb ua y 2, 2011. The genome p ojec
is deposi ed in he Genomes On-Line Da abase
(GOLD) [43] and he comple e genome sequence o
s ain MP5ACTX9T is deposi ed in GenBank
(CP002480.1). Table 2 p esen s he p ojec in o -
ma ion and i s associa ion wi h MIGS e sion 2.0
[44].
Figu e 2. Elec on mic og aph o G. und icola MP5ACTX9T
Rawa e al.
h p://s anda dsingenomics.o g 453
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 g un lib a y (GUIX),
a 454 Ti anium s anda d lib a y (GTWG, GWTA) and a pai ed end 454
(GSUN) 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 20×(454), 274X (Illumina)
MIGS 30 Assemble s Newble , VELVET, PHRAP
MIGS 32 Gene calling me hod P odigaL, GenePRIMP
Locus Tag
AciX9
Genbank ID CP002480.1
GenBank Da e o Release Feb ua y 2, 2011
GOLD ID Gc01833
BIOPROJECT PRJNA50551, PRJNA47621
P ojec ele ance
En i onmen al, Biogeochemical cycling o Ca bon, Bio echnological, GEBA
G ow h condi ions and genomic DNA ex ac ion
G. und icola MP5ACTX9T was cul i a ed on R2
medium as p e iously desc ibed [1]. Genomic
DNA (gDNA) o high sequencing quali y was iso-
la ed using a modi ied CTAB me hod and e alua -
ed acco ding o he Quali y Con ol (QC) guide-
lines p o ided by he DOE Join Genome Ins i u e
[45].
Genome sequencing and assembly
The inished genome o G. und icola MP5ACTX9T
(JGI ID 4088693) was gene a ed a he DOE Join
genome Ins i u e (JGI) using a combina ion o
Illumina [46] and 454 echnologies [47]. Fo his
genome we cons uc ed and sequenced an
Illumina GAii sho gun lib a y which gene a ed
42,620,699 eads o aling 3239 Mb, a 454 Ti ani-
um s anda d lib a y which gene a ed 146,119
eads and h ee pai ed end 454 lib a ies wi h an
a e age inse size o 9.3 kb which gene a ed
178,757 eads o aling 154.3 Mb o 454 da a. All
gene al aspec s o lib a y cons uc ion and se-
quencing pe o med a he JGI can be ound a he
JGI websi e [45]. 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 [48].
The 454 Newble consensus 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) [49]. The so wa e Consed [50]
was used in he inishing p ocess. The
Ph ed/Ph ap/Consed so wa e package [51] was
used o sequence assembly and quali y
assessmen in he subsequen inishing 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 [52] 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, un-
published) p ime walks. The inal assembly is
based on 29.1 Mb o 454 d a da a which p o-
ides an a e age 20× co e age o he genome and
975 Mb o Illumina d a da a which p o ides an
a e age 274× co e age o he genome.
Genome anno a ion
Genes we e iden i ied using P odigal [53] 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 [54].
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)
[55,56], 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 [57], RNAMMe [58], R am [59], TMHMM [60],
and signalP [61]. Addi ional gene p edic ion anal-
ysis 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 [62].

G anulicella und icola ype s ain MP5ACTX9T
454 S anda ds in Genomic Sciences
Genome p ope ies
The genome is 5,503,984 bp in size, which in-
cludes he 4,309,151 bp ch omosome and i e
plasmids pACIX901 (0.48 Mbp); pACIX902 (0.3
Mbp); pACIX903 (0.19 Mbp), pACIX904 (0.12
Mbp) and pACIX905 (0.12 Mbp), wi h a GC con en
o 59.9 mol%. The e a e 52 RNA genes (Figu es 3
and 4, and Table 3). O he 4,758 p edic ed genes,
4,706 a e p o ein-coding genes (CDSs) and 163
a e pseudogenes. O he o al CDSs, 68.8% ep e-
sen COG unc ional ca ego ies and 27.5% consis
o signal 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, and Table 5.
Table 3. Summa y o genome: one ch omosome and i e plasmids
Label
Size (Mb)
Topology INSDC iden i ie Re Seq ID
Ch omosome 4.3
ci cula CP002480.1 NC_015064.1
Plasmid pACIX901 0.48
ci cula CP002481.1 NC_015057.1
Plasmid pACIX902 0.3
ci cula CP002482.1 NC_015065.1
Plasmid pACIX903 0.19
ci cula CP002483.1 NC_015058.1
Plasmid pACIX904
0.12
ci cula
CP002484.1
NC_015059.1
Plasmid pACIX905
0.12
ci cula
CP002485.1
NC_015060.1
Table 4. Genome s a is ics.
A ibu e Value
% o To al
Genome size (bp) 5,503,984
100
DNA coding (bp)
4,759,459
86.5
DNA G+C (bp) 3,301,098
60.0
DNA sca olds
6
100
To al genes 4,757
100
P o ein coding genes 4,705
98.9
RNA genes 52
1.1
Pseudo genes 163
3. 4
Genes in in e nal clus e s
2 ,395
50.4
Genes wi h unc ion p edic ion
2 ,936
61.7
Genes assigned o COGs 3,259
68.5
Genes wi h P am domains 3,504
73.6
Genes wi h signal pep ides 652
13.7
Genes wi h ansmemb ane helices 1,108
23. 3
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 455
Figu e 3. Ci cula ep esen a ion o he ch omosome o G. und icola MP5ACTX9T displaying ele an genome ea-
u es. F om ou side o cen e ; Genes on o wa d s and (colo ed by COG ca ego ies), genes on e e se s and (col-
o ed by COG ca ego ies), RNA genes ( RNAs g een, RNAs ed, o he RNAs black), GC con en and GC skew.
G anulicella und icola ype s ain MP5ACTX9T
456 S anda ds in Genomic Sciences
Figu e 4. Ci cula ep esen a ion o he plasmids o G. und icola MP5ACTX9T 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. O de
and size om le o igh : pACIX901, 0.48 Mbp; pACIX902, 0.3 Mbp; pACIX903, 0.19 Mbp; pACIX904, 0.12
Mbp; pACIX905, 0.12 Mbp.
Table 5. Numbe o genes associa ed wi h gene al COG unc ional ca ego ies.
Code
Value
%age
Desc ip ion
J 160 4.45 T ansla ion, ibosomal s uc u e and biogenesis
A 2 0.06 RNA p ocessing and modi ica ion
K 249 6.93 T ansc ip ion
L
222
6.18
Replica ion, ecombina ion and epai
B 1 0.03 Ch oma in s uc u e and dynamics
D
33
0.92
Cell cycle con ol, Cell di ision, ch omosome pa i ioning
V 68 1.89 De ense mechanisms
T 212 5.9 Signal ansduc ion mechanisms
M 287 7.98 Cell wall/memb ane biogenesis
N
73
2.03
Cell mo ili y
U 123 3. 42 In acellula a icking and sec e ion
O
125
3. 48
Pos ansla ional modi ica ion, p o ein u no e , chape ones
C 174 4.84 Ene gy p oduc ion and con e sion
G 248 6.9 Ca bohyd a e anspo and me abolism
E 2 34 6.51 Amino acid anspo and me abolism
F
68
1.89
Nucleo ide anspo and me abolism
H 147 4.09 Coenzyme anspo and me abolism
I 126 3. 5 Lipid anspo and me abolism
P
1 37
3. 81
Ino ganic ion anspo and me abolism
Q 91 2.53 Seconda y me aboli es biosyn hesis, anspo and ca abolism
R
446
12.41
Gene al unc ion p edic ion only
S 370 10.29 Func ion unknown
-
1498
31.49
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 457
Discussion
G anulicella und icola MP5ACTX9T is a und a soil
s ain wi h a genome consis ing o a ci cula
ch omosome and i e mega plasmids anging in
size om 1.1 x 105 o 4.7 x 105 bp o a o al ge-
nome size o 5.5 Mbp. The G. und icola genome
also con ains close o wice as many pseudogenes
and a la ge numbe o mobile gene ic elemen s as
compa ed o G anulicella mallensis and Te igobus
saanensis, wo o he Acidobac e ia isola ed om
he same habi a [29]. A la ge numbe o genes
assigned o COG unc ional ca ego ies o
anspo and me abolism o ca bohyd a es
(6.9%) and amino acids (6.5%) and in ol ed in
cell en elope biogenesis (8%) and ansc ip ion
(6.9%) we e iden i ied. 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) amilies [63,64] in ol ed
in b eakdown, u iliza ion and biosyn hesis o ca -
bohyd a es. G. und icola 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 and D- ehalose.
Genome p edic ions o CDSs encoding o
enzymes such as cellulases, pec inases, algina e
lyases, ehalase and amylases a e in ag eemen
wi h biochemical ac i i ies in s ain MP5ACTX9T.
Howe e , he genome o G. und icola did con ain
many CDSs encoding o GH18 chi inases al hough
no chi inase ac i i y was de ec ed a e 10 day-
incuba ion wi h chi inazu e [29]. In addi ion, he
G. und icola genome con ained a clus e o genes
in close p oximi y 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 cellulose
biosyn hesis. We p e iously epo ed on a de-
ailed compa a i e genome analysis o G.
und icola MP5ACTX9T wi h o he Acidobac e ia
s ains o which inished genomes a e a ailable
[29]. The da a sugges s ha G. und icola is in-
ol ed in hyd olysis and u iliza ion o s o ed ca -
bohyd a es and biosyn hesis o
exopolysaccha ides om o ganic ma e and plan
based polyme s in he soil. The e o e, G.
und icola may be cen al o ca bon cycling p o-
cesses 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 Ac ademy o Finland and he New Je sey
Ag icul u al Expe imen S a ion.
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