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TitleMECHANISM OF POST-IRRADIATION DEGRADATION OF DNAIN A RADIOSENSITIVE ESCHERICHIA COLT NG30IRRADIATED WITH ULTRAVIOLET LIGHT
Author(s) Yonei, Shuji
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Issue Date
Text Version ETD
URL http://hdl.handle.net/11094/2153
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MECHANISM
sENsrTzvE
UGHM
OF POSM-:RRADXATION DEGRADAMZON
ESCM!MZCHIA COLZ NG50 IRRAD[AMED
Department
( Dectoral Mhesis )
SHUJ: YONE:
'of Biology, ?aculty of
Osa"ka University
OF DNA
wrmH
IN A RADZO-
ULMRAV:OLnv
Seience,
)fiCCHANISM OY POST-IRRAD!ATrON DEGRADAT!ON OF DRA :N A RADrOSENSr-
TrVE ESCHIIMZCHrA S29!Lll!ZZ NG50 rRRAD!ATED WITH ULTRAVrOI,]su! LrGHT
SHUJI! YONEI
DeparVment of IMoiogy, Raculty of Science,
06alua University
ZNff!RODUCEIrrON
An x-ray sensitive mutant, Escherichia coli NG50, iselated by ti')Kato and Kondo from strain Hlr30 is highly sensitive to ultraviolet
i) , and the phenomenon of extensive degradation ef eellularlight, too 2,3>DNA after ultragiolet irradiation has been reported. Kato arrd
3> have aiso found that the E. coU NG50 hsLs the follouring twoKondo --genetic characters ; A very lovr genetic recombiEation frequency in
the erosses vrith Hfr strain Bl of g. celi B sutd a very lovv rate ot
prophage ( e80 ) induction by ultravielet irradiation, as compared
with that of the vrild-type strain, g. sigtllLli !l/r50-R. From the
above bioche!nicaL and genetic analyses, the radioseEsitive S. sg,1!ÅÄ,i
(1)
NG50 was classified phenotypically as a mutant belonging to the ReeAe
type in the three types of reeombination-deficient mutants of E. s;2,1-,i
4, g)K12.
ReeA mutants of E. eoii KZ2, as well as NG50, are also highly
4-8) 'sensitive to uitraviolet iight and shows an extensive degradatiorr ofceiiuiar DNA by uitravioiet irradiatioEiZ'8-i2)rt is sure that there is
a close relationship between the muked ultrasztoiet sensitivity and
the distinctive nature with extensive DNA degradatien by ultraviolet
irnadiation in the Re(A' mutants.
9) Ciark, Chamberlin, Boyae and HowardeElanders, a!zd Howard-Fianders
. 13)aila Boyce pestulated that the post-irradiation degradatiou of DNA
in RecA- mutants would be induced by the single-strand gaps in DKA 'nrolecule6 vvhich were produced by incomplete dark repair events.
iD, "> Eerii and Suzuki showed recently that the ultravLelet-induced
DNA degradation of a RecAe mutant of E. eoli K12, Je 1569b, occurs
on a speciai DNA pertion involving DNA replicating poirFt, because
the pulse-labeled DNA regiolt begins to degradate prior to the deg-•
radation of pulse-labeled and chased DNA region in growing cells.
'rhey suggested that the replicating point is a sensitive site for
trradiation, because the dark repair of pyriutdiEe diners or the
(2)
repMaa•tion of DNA nNist be eontreled by certain lltechanism in order
that tke replicat LBg point does Rot run across' the site under the
repatr.
4.o, iS) The experimental results and speculatiolts by the above authors
might mean that the DNA portion far trom the replieating point
would not begin to degradate until the replicatiing point' reaches
to the pyriutdine dimers on the DKA pertiort, or uiseU the degrada-
tion initially occured on the replicating point comes aleng the DNA
'strand to the damage ot the DNA porcton.
Hewever, we have observed' preliminary that g. sE2,1-i NG50 u!rder
the conditions considered to be absent or DNA replication is also
very sensitive to ultraviolet irradiation and a marked degradatien
of ceilular DNA occurs, too, aften ultravioiet irradiation.
Aithough the rsechansm of degradation of the chronsosome invelving
replicating point llrast net be the same to that of the chromosome
without D"A repUcatien, it is very interestiBg to investigate the
degradation of DNA by ultravi.olet irradiation of the ceils yvithout
DKA replication, especially to make clear the eorrelation between the
fragmentation or degradation of DNA molecules and the early events
operattng to the pyrimidine diners on DNA mole{rules.
(3)
The present experimentai results will make evident that, there
is an extensive post-irradiatiorr degradation of DNA in the ehromosome,
too, not involving the replicating point and the initial cause for
the degradation is the single strand gap resulted from the excision
of pyriutdine dimers whLch is the first step of dark repair process
for ultravLolet damage en DNA molecules.
bKATERrALS AND METHODS
Bacterial strains and media
The bacterial strains used in the present experiments were g.
coli NG50 ( recAuvrt ), NG50ELMvgrC ( recAuvre, double mutant ) and
the wild-type strain, H!r50-R ( rectuvrt ). All of them were argi-
3)nine-requiring imitants. They were kindly provided by Drs. T. Kato
and S. Kondo in the Department of Eundamentai Radiology, School of
Medicine, Osaka University. The cells were grova at 570C with
aeration by shaking in )E9 medium supplemented with 2.5 mg/ml of
Ditco casamino acids ( denoted as E"E9 ). Tbe M9 medium coEtainect
the following irBgredients per 1,OOO uut of water ( pH 7.2 ) : 1.1 %
(4)
Na2HP04, O.5 % KII2P04, Q.5 % NaCZ, O.i % Nff4CX, O.4 % glucose. 20 al
O.OZ M CaC12 and 10 ml O.1 MMgS04. 0ne geEeratioli time of g. sigtgU
NG50 "as about 60 min in EN9 mecM.urE at 57PC.
U].traNxtolet irradiatLoE
' The suspension of becterLa was spread in a Petri dish to a
aepth of 1 ma and irradtLated with a gernrtcidal 1anp ( Mitsubishi GKL-
iO, 10 W ) with gentle shakittg at room temperature. The concent-
racaon oÅí baateria Åíor trradiation was usually about 6 x lo7 ee11slal.
The dosG rates used we]ro 2 ergs/nm4/seer and 2o engs/mm2!seq. irhe
intensigy ot ultrautolet lignt was determuted ntth a Meshiba UV phetoe-
meter ( Toshiba Elect. Co., er-1, Japan= ), which was correeted by
Dr. Laterjetts UV-photometer.
Anal sis of DN s thesis of bacteria uuder various conditions
cells were grown in ]gMg mediuur to approxtmately s x lo8 cens/ml,.
'washed twice and resuspended in five times the voiusse of M9 medium.
rhe eultureg in 5 rniL were then suppiemented with 5 pC/5pglrnl er
[HZ -thymidine and 200 ug/ml oÅí deoxyadenosine. in order to pro-
vide two kinds of cell conditions, groyvth condition and amino acid-
(5)
starved goB•ditien, they were divLded into t.wo portions and then incu-
beted at 579C with oir vvithout 2.S mglul oÅí easawtno acids. At
regular int'eruaZs, O.2 ml of the eultures was taketr into the ice-
cold 5• % tstehloroaeetic acid' ( TeA >. fhe aeLd"insoiuttle fractione
were collected en glass fiber filters ( Toyo GB 100, Japan ), washed
and dried. The radioaettvity of the fraction was eounted using a
Uquid ScintiUation Counter ( NRcitear Chicago Mark Z, U.S.A. ) in a
toluene-DPO•-POPOP ut.xture. In the experiments ot DNA synthesis
tn the presenee of ehleranphenicol ( CP( ), tihe beLcterial eells were
incubated ivith theVHJI]-thyutda"ne at 579c in the presence of cM.
XL!SWs2!!R!uti-gsnt.IA!us-slgga:swesu!tDNAdrdti
ltL:.-SXg"sc!2z.-{lf2A,s!=2,!IEgiL3"N,s}-.IEgSs2Ea,g],An Elisbcidlnsolublematepils
The Neasurement ot post.irradiat ton degradation ot DNA by the
change ef anount ot acid.insoiuble materials in the cells under a
growth condition was carried out as follovs ; ]3acterial cells cultured
overnight were diluted forty rold with the fresh EM9 medium su?"le-
mentea vvith 5.5 pC/3 pglul ot[Hb-thymidimo and 200 pg/tl of deoxy-
adenoBine. The eell suspension. was incubstted aerobically fer 2 hr
at 570c. The cells were harvested, washed three times, resuspended
(6)
iE tive tin}es velume of the E"E9 medium and irradiated wtth different
doses of ultrauSolet light at room temperature. rhe non.labeied
thymi(bee was added to the irradtated eultures in a eoncentration
or iOO pg/ml, euid O.2 ul vas withdrawn Åírom the cultures into the
lce-•colct 5 % MCA at reguiar interuals threugh the post-irradiatien
incubation at 5?PC. Mhe acid-insoluble fraetions were eoliected
on glass-fiber fiiters, diriect and counted using a Liguid ScLntUlation
Counter. 'lhe degree of DNA degradation in the eells through the
post-irradiiatien taeubation' was indicated as per cent of radieactive
eounts in the acid-inselubie fractions of the ceUs withdrawn
irnmediately after irradiation without incubatien.
The measurement of DNA degradation in the antao acid-starved
'cens was carried out as tollews ;[H5J .thymidine-labeled cells in
expenential growth phase vrere washed twice vith M9 medium containing
ne arimo acid, resuspended with five times volume ot the M9 and
incubated at 570C fer 90 ntn tto starste the ceils from utno acids.
The amino acid.starved cells vere irradiated with difterent doses
of ultranelet Ught. The experimental procedures after the irr-
adiation were the smu as aescrlbed abeve.
in the experiments for DNA d'egradation of CM-•treated eells, the
(7)
vvashed cells labeled ini.th[H5]-thyntdine were incubated at 570'c fer
90 min in EPf9 vrith 50 pglul eÅí CM and then irradiated wtth ultra-
violet light. ' The cells with DNA molecules labeied yvith[H5)-•thymidine on
their differeEt PXA region were prepared. The cells having DNA
strandis labeled only near the in-itiation point of DNA repUcatiott
including the point vere called as the cells eontaining labeZed
t, early region ,' of onA. The gells havtng DNA labeled eniy tar
region frorE the inttiatLen peint of DNA replicetieR were calZed as
the cells containing labeled ,t late region ,i ot MIA.' The fonv
celis were prepared as follows ; The eeZls Btarved amino aeids tor
90 utn were 6upplemertted ntth 2.5 mglml ot easantuo acids, 10 "e/iEZ
of[H3]-thymidine and 200 pg/ml of deoxyade"osirte bo allow to initiate
the DKA replication. Atter incubation at 570C for 2e min, the
labei was ehased wtth a 1arge amount of non-labeled thyrnidine and
tlae aells wer6 incubeted for more 80 nrtk ntth 50 pglul. :n order
to prepare the latter, the arnino aeLd-star"ed cells were supplets-
meRted first with casantno acids alone ut incubeted at 570C for 20
utn, and then [H5]-thymidine, deoxyadenesine and CM were added to the
aeXl suspension iu the ceneentration of Ze Fe/ul, 200 pg/al and
(8)
50 pglmL, respectively. 'rhen the cells were incnhated for 80
utn at 570C. The experimental methods Åíor the measurement of
post--irra(!iation degradatiett oÅí DNA in the ceZls iabeled ta difter-
ent DNA regton were essentialZy the sane as described above.
2. Sedimentation anal sis b alkaliite sucrose-densit radient
The post-irradiation degradatien of DNA was also analyzed by
sedimentation teahniques. The sedimentation analysis of DNA nxole-
eules in al}ta1ine suerose gradient was cetrried out essentially in
14>the sane metbocts as described by McGrath and wrIUans. The cells
were harvested by a centrifugation at 6,OOO rev./min Åíor 5 min
and resuspended in O.Ol H Tris-ttCl buffer ( pH 8.1 ) coutaining '14 % sucrose and O.Ol M EDTA. The eell suspension was incubated
at 570c tor 5 min with Z mglml of lysozyime to cenvert the cells
into the spheroplasts. The spheroplasts in O.l al of suspension
vrere placed on the layer of O.5 N NaOH ( O.2 rEl ) whieh had beert
layered on a linear gradient oÅí sucrese ( 5- 20 % in 4.4 ml )
dissolved in O.2 N KaOH, O.7 M NaCl and O.OOI N EurA. Usually,
about s x lo6 spheroplasts were placed on the gradient. The
sanples were centrituged for 90 atn at 50,OOO rev./!nin at 40c in
(9)
a swinging becket roter ( RPS-40 ) of Hitaclti Preparatime Ultra-
eentrifuge ( sTapan ). After the centrifugatierr, two-drop tracstorrs
vere eoliected trom the bottom of the centrituge tube piereed' ntth
a thin needLe on pieces of filter paper ( T`oyo No. 7 ). The pieces
vere suscessively washed with 5 % TeA, 9S % etlianol and ether, and
then dried. RadieactLwtty oE "he paper was countect using a Liquid
Seintillation• Counter in a toluene-DPO-POPOP mixture.
:M:s2gs-xs2}ug!us-2z-!g,s-sg.-gs!gcee ntof ilai{MtrdLrs
Measurement ot. pyri!nidine diners in DNA tuoleeules was carnied
eut a6 feUovs ;[H5J-thyrliidtne-labeled cellB were irradiated with
ultrantolet iig}rt. The eells "ere precipitated and washed' tvice
vith 5 % reCA. The precipitates vere the"' vashed vith iO mL each
ot 80 % ethanol, 99 % ethanel artd a mixture of ethanol and ether
( i : 1 ), suecesstvely by cert-trifugatien. The yvashed precipitates
"ere utspersed in 1 ul ot S. % TCA ut incubeted in a vater bath at
950C tor 50 Nin. fEhe extraets containtng labeled I"IA ivere treated
vith ether repeatedly three times to remove t'he Ton trom the extracts
and then coneentrated to about O.l mi by eveporation. The DKA
selutien was hydrolyzed ntth 99 % forntc acid' for 15 min at 175 -
( ro )
18eOc in a sealed giass tube sgLturated with nitregen gas. The
'hydrolysates were applied on a paper strip ( Teyo No. 51, 2 X 40
cm ) and deveioped vdth a mixture of n•-butanol-acetie acid.water i'ty)C 80 : 12 : 50 ) at reom temperature by ascending chrematography.
In the preBent system for chromatography, the Rf values of thymine,
thymine dimer and thyrni.ne-uracil dimer were O.6, O.5 and O.2,
respectively.
The measurement. e'f pyrirnidine dimers released from the irra-
diated eeils into acid-soluble fraction during posteirradiation
incubation was also carried out by esseEtiaUy the same procedures
as described above. Materials in the acid-soLub e fractions
were hydroiyzed with tormig acid and chromatographed by paper.
The amounts of iabeied "ueleotides and oiigonucleotides ineiud-
ing pyrinddine diHters in the aeid.soluale rractions extracted from 'irradiated eells were meagured. The aeid.extraeted materials•
were eharged into Dowex-i ( C3.' type, X 8, aOO e 400 Eesh ),colurur!
( 1 X 2.5 em ) and eluted stepvise with water and then with O.5 N
HCi. Labeled nucleosides are eluted by the former step and all
ot the labeled mucleotides are eluted by the latter step.
(11)
Other remarks
' Lysozyme ( E. e. 5. 2. 1. 17 ) was purchased
co. Ltd. ( u.s.A. ). [H5]-thymidine vvas obtalnd
active Isotope Corporation ( !okyo ).
from Sigrna Chemical
Åíront Japan Radio-
Dose-survi.val. curves
RESULTS
of S. coli H r O-R and KG O irradiated vvith
Nl,9zexi!,stEte32,a,aa{!Ltviit1ht
Cells of g. sEgt,-,li Hlr50-R and NG30 in groyvth phase, 6tationary
phase, starved con(tition for utno acids or a condition pre-incubated
with CM were irradiated with ultraviolet light, respectively, and
the colony forming abilities were examined. The numbers of coiony
were plotted agatnst ultravLolet dose used in ?ig. 1.
( stg. 1 )
:t was shown that g. ggtA,i!,U NG5e, eygen if the cells were in differ-
ent physiological conditiens, was still highly sensitive to ultra-
( 12 )
vioiet light as eompared vvith the wiid-type strain H/r30-R, although
the a. s22,1-,i NG50 in stationary phase, amino acid-starvation condition
or pre-incubated eondition with CM was a littZe less sensitive in
every case than the cells in growing phase.
De radation et DNA of E. eoU NGO in rowin hase after uitraviolet
irra(liation
The mode of DKA degradation in grovvirrg phase of S. sigt,-li NG50
'after uitraviolet irradiation was inves"gated. The ceils, oÅí 'which DNA nvas uniformally labeied vith[H5)-titymidine, were washed
and resuspended in EM9 medium. The eulteres vere inerrbated at 570C
in the same medium after ultraviolet irradiation. The doses used
were 2o, loO and 400 ergslmm2. Tbe reuiaining radioactivLty in the
acid-insolub].e fractions was deterntned as described in Materials
and Methods. The results are shovvzi in Fig. 2.
( st g. 2 )
:t was shown that the radioactiutby in the aeid-insoluhle trac-
tions was rapidly deereased during the post-irradiation incubation
(15)
in each case. Howevere it was also tmown that there was rto linear
relationship 1eetweett the rate or. deg.radatiok and tke uXtraviolet. dose,
and vrhen the eeZls in grewing phase were irradiated. even enZy by a
sma11 dose of 20 ergs/an2 resuited an extensive degradatlen of mgA.
A marked spontaneous degradation ot DNA ot unlrradiated eslls was
also observed.
Sto aeof' DN'A re lieatien • wtno acid-starwation• and b chioram-
2ahg-ggsenico1
rhe DNA replication durikg utno aeid-•starsratiou was examlned
through the incorporation otLH35]-thymLdine inte acid-insoiubie Åírac-
tions. Fig. 5 sbows that the rate oÅí incorporatieE oÅí the labei
( Rig. 3 a, b)
in the utno acid-starved cenditions are slowly deereased initiaily
vvith the tirae ef ineubation and finaliy aompietely stopped by abeut
9e min tacubatien in beth strains.
The restoration of DNA repiieation was alse shewR in Fie. j5b.
When the starved cells, of NG50 were transtered from the M9 mediunt
(14)
bo the E"f9 nrectum, the rate ot incorporation of label rose quiekly
te nearly the sane rate ef DNA replieation' oÅí tbe cells supplementÅëd'
with arnino acids. ' , Fig. 4 shews that the chloramphenieol steps eompletely the DNA
replication ef the ce11s} E. sEgtsill NG30, evetr if the aatno acids were
atrandantly present in the medium, by the incubation wit'h the drug
Åíer about 60 e 70 utn as well as the case ef amino acid-starvation.
'
( Xig. 4 )i
De radatien oÅí DNA or•E. eoli NG e after ultraviolet irxadiatien in
3thigR-SSEESt!ISiS2-.9S-ll!tSL!:W,gsu,s21hS lefDNAreliCtion
a. eorrelatien laetween de radation rate andi ultraviolet dose
The,degradation ef DNA ef E. ssul,iLi NG50 aft•er, ultravielet irra-
diation in the absence oÅí DKA replica•tien was exautned. rehe cells
were• starvedi• amine aÅëid$ by the ineubation fer 90 isi.n in "G9 rEedium
'and then' irnadiated with different doses of uitrwtolet iight.
The mode of the loss' et radioactivtty from tlae a"Ld-insoluble frac-
tioEs in tbe ceurse ef time ineubation at 57QC iii )e9 medium are
'shown in Fig. 5.
(IS)
( ?ig. 5 a, b) .t
' '
:t is ebvteus in Yig. 5a that the DKA of the cell$, E. eoM KG50 " --e--b
in which any DNA replication has been eeased is also extensively
degradated as well as the case ot cells iri grovTirtg phase, although
the degradation rate, espeeially at the low doses, is smaller than
that in growikg cells having repUcating,point ( Reffer Mg. 2 ).
The spen-taneous degradation of DNA is alse less in the starved ceUs
than the grewing celis.
tievever, the rate oÅí DNA degradatien in this physiologieal state
of NG50 seerned to eorreBpend more well to the ultraviolet dose than
'that in growing cells.
rhe results in IFIg. 5b were shown' as retereuce data. The
post-irradiation degradation of DNA in utno aeid-starved E. sgt,-,li
Elr30-R and S. celi Bs.1 were shown. Oniy a linttted degradation
srras observed in g. sEgo,-,li Elr50-R irradiated wLth 1,oOO ergs/rnm2.
It is sure that the phenomena are related to tlxe alxLlity oS excision-
repatr systems tunctioutng on the uitraviolet damage to DNA rEole-
eules, sikce g. sEgt-,li Bs-1 which lacks the abi.lity to exeise pyrie
(16)
iG)ntdine dimers exhibits no
' The DNA degradation
S. coli NG50 of whieh DNA
incubation for 90 ntn iR
DNA degradation.
during the post-irra(utation ineubation of
replieation had been stopped by the pre--
the presence ot CM is shown in IMg. 6.
( Eig. 6 )
rt was also shown
dorm in course of tine
starved cells. The
ultrawtelet doses.
that the DNA ot E. ggts},li NG50 broke rapidly
Qf inqubation as the ease of amino acid-
rates oÅí degradation eorrelated welX te the
b. Com .ar-son of ost-•irradiation de radatien ef DNA in E. coM KG O
]Z,gJlsll!,gs-!L.sl2,i"igxs!ES-l2!llL.!gat,stlygb1didiff tDNAr
rhe experinents were designed to stud'y whether er not the post.
irradiatiott degradation of DNA might begin at a some specitie point
even it the DNA had no replLcating poirEt. Two kinds of cells
of E. coli NG50 labeled in different MgA region ( ,, early regLon ,, ' - 'and ti late region tt ; see )fateri.al.s aka Methods ) vere prepared.
' 'rhe loss of radioactivtty from the acid-insoluk[Le fraetions was
( 17 )
respectively deterulrred• in these two kinds ef eeUs, and the rates.
of pest-iruadiation deg.radation eÅí labeled DNA region were aompared
ea(th other. rhe results were shoyvrE in ira.g.. 7.
(+ reig. 7 a, b. ).
' Mhe quite analegous results were fouad in' the both experim'ents.
The rates oÅí post-irradiatien ot DNA by the sane uitravioiet dose
were r!eayly equal to each other in bpt.h DNA region•s. These results
indicate that the pest-irradiatien degradation oÅí IXYA oÅí g. sEgte,a,li
NG50 in the absence of DNA replication does rxot be induÅëed at a spe-
gitie site in osA molecules but can begin at many sites of DNA at the
seLrke time.
c. ,De radatien of DNA of E. eoli NG OuvrC after ultravLelet irradia-
rn order to know the contribution ef diiger-exeising phenomena
to the pest-irradiation degradatien of ceUulatr DNA, the degradation
of DNA of a double mutant in recA aitd uvrC was exarntned'. E. eeli - --"-e-ep--pa' ,' , ts!eere' whiclr is lacklng or iealry in the abUity of diraer exeision
(18)
was used. rehe experimental result.s oÅí IMA degradat'ion during post-
irradiatien ineubation of tbe eelis, irradiated wLth. verious ultra-
violet doses vvere shomi' in Rig. 8.
( Rig. 8 )
rt was found that the post-irradiatien DNA degradation oÅí S. sgtsli
NG50 was markedly suppressed by the additional mutatiou oÅí s:tyzC.
However, it was also known that a sig!rifica!rt degradation'• of DNA was 'still ocaured in the eells ef g. s2,iE,isi NG50uvre.
The araeunts of dimers in the DNA of the cells incubated for 60
and 120 min after ultraviolet irradiation of 4oo ergs/ua2 were
iaeasurect, respevtively. Eig. 9a shews the paperehroruatograrns indi-
caging the time-dependent decrease oÅí the amount of pyriutdine dimers
during post-ireadiatien incubatieu oÅí !. sg,1-i NG30. The resuZts
'by !. sift2,-li KG30LvrC' were show!t in Rig. 9ke.
( Ri g. 9 a, b )
A decrease oÅí pyrirr tdine dimers from the DNA in accordance with
( 19 )
the
but
were
d
time of post-irradLation
no removaL of pyrizniutne
significantZy deteetedi.
tteet ot e ffeine orr the
incubation was apparent
diners from. the DNA oÅí
DK de radatien in E.
in
E.-
E. coii NG50, --set,i,i!,lLNG:5QLvrC
croli GO
The erfect ef caffeine on the onA degradation• in the eells of
S. wrii NG50 following ultravielet irradiation was studied. Caffeine
was added te the cultures immediately betore irradiatien• of 400
ergs/an2. mhe results are slhown in' Eig. 10.
( ys.g. io )
fl!he degradation of onA of S. sgtsl,li NG50 during post-irradiation
incubation, was inhibited in accordanee"•vith the increased concent.
ration of caffeine.
rt was conTfirmed Åírom the above alZ experimental result6 that
the radioseissitive becteria, !. sgti,;!,li NG50, shevr a rnarked post-irya-
dLatlon degradation of DNA egen if tlve cssXls vrere noti in growing
phase but in a resting condition in yorhich there is no DNA repliaa-
C 2Q )
tion. In addition, the above experimeatal results aUowed also us
te assume that theepost-irradiatien degradation of asA must be eaused
by the exc Lsion oS pyrinddiEe dimers producea in ulptatlolet-irra-
diated onA.
If the degradation ef DNA of S. gsul,2,i NG30 was directly related
to the excision oÅí pyrimdiue diiiters, fragmeEtatien oÅí DNA shouid be
accompanied with the release of dinters Åíeota tire DNA, which produees
single-strand gaps in DNA strands, foUowed by progressive degrada-
tion oÅí DNA moleeuies.
AnaL ais of de radatioR h seclimentation rotiies oÅí bNA in aLkaline
EtsgsgssLsXs}iNi,usmaga,g!x!iucredxt dient
The patterns oÅí DNA in the ceUs of g. Eg,Zgl,i NG5e in a eondition
yvitheut DNA repliaation were analyzed in an aikaUne sucrose gradient
in relation te the post-irmadiation dagradation of DNA depending to
the ultravioiet doses and/or to the time ot post-irradiation ineuba-
tlen.
a. Znitial fr mentatiorr ef DNA b exetsieit of rintdtse dimers
The amino acid-starved eeils labeled with[H5]•-thyniidin'e were
( 2i )
prepared. Ihe ceUs irradiated with 2oo and 1,ooo ergslmm2
respectively, and allewed to incubate only Åíor 5 ixin in M9 medium
at 37eC were harvested' in ice.aora' terRperature, 1ysed by lysozyme
and eentrifuged in alkaUne sucrose.density gradient. The sedi-
mentatiolt profiles were shova in Fig. il.
( Mg. Il a, b)
It was seen that the main peak oÅí DNA distribution was shifted
more to the smaXler size eÅí DNA in correspondence to the increa$e
oÅí ultraviolet dose even ilt the radieresistant cells oÅí Hlrj50-R. 'the shifting rates of the DNA peak trom the peak oÅí unirradiated
cells vrere nearly the sane each other in both strains.
The facts indicate stroza'giy that the Lnitial events fellowed
by extensive degradation' of DNA in g. s}suisi NG30 during the post.
irsadiation incubatien is the removal of pyrimidme dLmers frem the
irradiated DNA retwtlting in single strand gaps in the DNA meieeules.
b. Post"rradiation de radation oÅí DNA ta S. coXi RG e in corres-
12sesssg!!si:Bdencetoultravioletdoses
(' 22 }
Mhe utne aeid-starved E. eeli NG30 ceUs were irradiated with - --y 'ditferent doses ef ultranolet ligne. The eells were then incuha-
ted for 2e min at 570C in the M9 Ee(lium. Tbe patterns ef the DNA
oÅí the eells wolre eompared ts reiation to the ultravLolet dose.
rhe results wera shovm in Eig. 12.
( Fig. i2 >
rt is ]snowa thae tbe wtrr peak of asA distrikriitien was shiÅíted
more to the sma11•-size ot ERA iu correspodeBee to the increase oÅí
ultrav:.olet close, and that the distril;ution of DNA beeomes hread as
compared with that of non-irrautated celZs.
The zaeeut size of eaeth DNA distritntien shown in Yig. I2 was
compared with thGtt of mon-irradiated eelLs.. The reXative values
of number average moleculsLr rveight of DNA of the eaeh dLstribetion
irr )were calcuiated using Kapp and Sptthts formula assuming the rEole-
eular weight of DNA from non--irradiated cells as 1. Ihe values
were O.72, O.57, OJ7 and O.26, for 100, 200, 400 and 1,Ooo ergpus/mae
respeetival.y.
( 23 >
c. De adatieii. of DNA in E. eoU NG/, durin st-irradiation incu-
bation
S. sEsi,1-.i NG30 and H/r30-R were ttsed. Mhs c'alls starved uto
2acids were irradiated with 200 or 1,OOO ergs/an and then incubeted
at 57ae ÅíQr 2e and l20 atn in the M9 medium. rhe lysed eeils were
centriÅíuged as beÅíore. Mhe sedtaentatiort profUes sLre shown in
Yig. 15.
(' Fig. 15 a, b. g• >
:'t was found that the degradation of DNA in the irradiated
E. eeli KG3.0 progressed okly into shorter fragmeut of DNA in eerres--pondnece to the tine eourse oÅí post"rradiation ineubetiort. <1 Rig.
I5 b anct e > Both t.he moiecular size af DNA strandi' and the aedct-
precxi.pitable amounts oÅí DNA at the post-•irnactiation incuhatioEt of
120 ntn are s!uatXer tharr these at the time or. 20 miu'. rhis pisemo-
meEolt is more distinet in the eells: irradiated wtth 1,ooo ergs/ua2
than the eells ntth 2oo ergs/nn2.
On the contraTy, in g. sg,71! L,ii H/'r50-R ( rvig. i5 c ), it was tmomi,
that, althougtT the irradiation with 1,OOO ergs/ua2 resulted in a
(24)
smaU size of DKA strands in alknline cortdition at 5 min incubationt
the muA moiecules did not ctoEtirEue to break dewE' inte sua11 pieces
but/ recrexnerea almost to the moleÅëuistr stze oÅí IXgA in nen-irradiated
cells during the post-irradiation Å}neubation ter 2e ntn.
The change in molecular size of DNA in g. eoZi NG50 in course
oÅí time ot pestt.irractiettion incubetion was summarized in rable r.
The moieeuiar size of the main peak in DNA distriXnition was represented
as a distance frour the menisws in alkaZine sucro$e. gradient.
( TabLe r )
Release of rintdine dimers frem' irradiated E. eoli NG O into aeid•'-
selubie fraction
rn erder to make sure the hypothests that the initiai eveEt
for the induetion of' DNA degradstion in uZtravieiet"rradiated E. coli
NG350 is the fragmentation of DNA eaused by remeva1 ef pyrimidine
dtuers Åírom DNA moieeuZes, the cells were irradiated with various 'doses of. ultravielet light att'd the labeled nucieotides in- the aciG-
soluble fraction extracted trom the eeZls incub&tedi f'er 5• vain• after
ultraviolet irradiation were measurect• by eoluma ehromatographic., tecÅ}ir-
(2S)
nigues. Ihe amounts ot- label were plotted against ultraviolet
dose in Eig. I4.
( Rig• 14 )
rt is clear that the removai of pyrimi(ti.lte dimers from the
ultraviolet-irpadiated onA oecurs' as an eariy evez!t for the post-
irradiation degradation of DNA of S. sEt,lt,si KG50, and that ehe release
of dinters is iiiore ln acaordance with the increase of ultravioiet
dose.
The released' pyrirnj.dLne dimers were detected by papnr-chromato-
graphy in the acid-soluble fractiens extracted trom the cells incu•-
bated for S rnin aÅíter ultraviolet irradiation by a dose of 2,220
ergs/mrlt2. The results were shown in Mg. 15.
< Fig• 15 )
rt is apparut tnat the acidi-soluhle fractions of S. ggt,sELt: NG]50
'contatns pyrimLdine (iimers exaisee' from the onA, even when tbe
fractions were prepared Åírom the eells incubated fer only 5 mS.n
(26)
after ultraviolet irradiation.
Frem the above all experimentai results, it ean be cencluded
that the extensive post-irradiation degradatlon ot onA in E. colL -t- -NG30 is induced by a progressive DNA degradation expanding indipendent-
ly from completing the repair oÅí single-strand gaps in DNA nrolecules
preducea by removai ot pyrimidne dimors.
Drscuss:oN
The present experiments cieared that a recombination-deficient
rautant, g. ggtskli NG50, is very sensitive to ultravioiet irradiation
and a degradation of ceilular DNA occurs extenatveiy aÅíter ultrats
vi.olet irradiation, even vken there is no DNA replieation iri the
cells.
B) MaalÅëe and Hanawalt have reported that, aithough atnd.no acid-
starvation allovs to complete the present cycle of DNA repUcatien
to the starved eells, it never allows to initiate the rtext eycle
ot DNA replication. It is apparertt, from the experimental resuits
( 27 )
about the ineorporation of[H5]-tltyintdine inte acid-ingolubLe materials,
that the present experimevatal eonditions, that is, tacubating the cells
in M9 medium er incuhation with caloranphenicol, hleek completeiy
te reinitiate the nevv crycie oS DNA replication in S. sigt-,ii NG5e.
The phenomenon that the DNA molecuies ef Z.• eoli NG50 in whieh
there is no onA replication were aZso extensively breketr down by
post-irradiatiort ineubatien as in the growing cells in which the DNA
continues to repUcate was confirmed by tue rapid deerease in the
amount of acid-precipitable labels' and by the conversion of inttiaZ
cellular DNA to smaU fragittents oÅí DNA, aithough the rates of deg- 'radation of DNA in the growing. eells were much greater than that in
'the resting eells wkett the ceXls were irradiated with reiatively
iow doses ef ultraviolet light.
Moreosre?, it is aiso evident that the degradation of DNA in the
resstng celis of- g. si2t,-ii KG30 does not begin at a speeiÅíic site of
DNA, tut can occur from !urmy sites of1NA at the sane time and at tke
seave rate in invadiated eeZls. '
The assumptLo]t that the first even•ts for the induetion of pest-
irradiation degradation of DNA havtrtg no repUcating potats iEust be
the exÅëision oÅí pyrimidine dimers frorza the DKA is reasomably accept-
(28)
abLe by the foUoveing two facts, <1) the DNA moLecules in the
irradiated eells of S. ggtkiliE NG30 incubated fer only 5 nhm aÅíter
ultraviolet irradiation has beetr eut inte smaller Åíragmerts as welZ
as that observect in the wUd--type strain .Hlr30-R ;<2) there is
'aetually the release of pyrintdine dimers tato acid-soluble fractioEs
frora the eeUs ine.ubated Åíor the shert tinc aÅíter ultraviolet i;ya-
diation. Mhe assumption was further supported by the facts that
the post"rradiation degradatien of MYA was markectLy suppressed in
a double mutant, E. sist2,EilL NG30uvre, arid inhibited by the presence ef
eaÅíteÅ}ne. :t is known that caffeine imhibits tke excision' of sg,2n )pyrimidine dimers by Åíorutng a complex with ultravLoiet-•irradiated
i9 ) The residual degradation of DNA in irradiated eeils oÅíEMA.
E. coli NG30uvrC raay be due to a low aetivity oÅí dimer-excision- M)detectable in irttadtated' uvrC mut'an'ts. ts Zt is sure that the DNA moleeules in growing cells of g. eoU
NG50 ifireBe IMroken extensiveZy down even when the ceUs were irradiated
ntth only a small dose ot aO ergs/rllma, as the ebserwations by Horii
g]t; g"i.iO)on a ReeA- mutant of g. sigll,i!,i K12, Je l569b, atthough -E- sg,ll,si
NG50 in resting state shows no so gfeat po6t-irradiation degradatiolt
ef DNA by sueh small deses.
( 29 )
Zt may be true that a different mechanisit must operate on the
degradatiolt process of DNA in growing eeils from that in the resti.!gs
cells.
The author would like to speculate about the distÅ}ntrt phemomena
in growtng ceils as that, the enhaneed post-irradLatien degradatien
of onA may be abesulteet frora sueh a syRergistic eÅíÅíeet as that the
functLon of onA replication disturbs furthermere the proeess ot DNA
degradation indueed by a predoulrtrant degradation oÅí' DNA uneoupled
well to the effective repalr oÅí ultrautolet iesions, through runRing
oÅí the replicatlkg point6 across the degradatien sites in msA strands.
The above speculatiort- means that the initiai e'ause Åíor the post-
irradiat'ion degradation of bNA in E. sigtÅíli NG30 is dtmey-excisien,
and the produaed gaps would be enlarged by a predoutnant degradatiorr
event Rot eo-ordinated weil with the process' of repalring the gaps.
The arrival oÅí vNA replicating po' ints to such the diserdered'.degrada-
tion site weuld raake the egentEsL iiiore compiex occuring to the deg.rada-
tion sites, resuiting in drastie DNA degradation.
") - There isareport thataReeA unitant ofg. sst-,li M2 casmecover
significantly,trom ultuaviolet-inactivatien by ineubating the irra-
dtated cells in buÅíÅíer soiutions. The phenomana et iiguid hoiding
(' 50. )
reaoNFery. wouZd be relat:ed t'o the pr.esent'r experi'merrkaliacts tlta't
tbe post.irradiatiorr deg.radatiorr' oÅí Mlll ts, autmo acid"st.arwedi
Spt sigtiIE,i KGE50 vas geeatly' swppressed by entttin'g glirco$e frerr tbe
NE9 meetuif e See,. n- g.. ts )..
( Eig. 16 )
Anyhow, a phy"otypical character oÅí S. sstl-,U NG50, high sensi-
tivLty eo ultraviolet irradiation, imi$t be essentiaUy related to
tbe phenomna observect at the irradiated ceXlular DKA whiek beealsts"
extenatvely dovvn by pest-irradiation incubation.
srmoRy
rehe pest-irradiS.atioE ctegraaa•tioE of DNA
mutant, g. ssl,1!si KG50. was irvestigated. An
of DNA eerrespe!TdiHg te tbe ultravi.o]Let dose
eeUs vrhieh rep].Leate Ro msA.
The post-irrad'iation degradation ot DNA
(N)
ta a radXoseE-sitive
' exte!tstve ctegradation
was' observeditr the
having no replicaging
poMts does Rot begin to oacur fr'om a specific point of msA, but
do oeeur tO mEmy sites of irractiated osA at the same tiite akd' deve-
iops at tne same rate.
rhe tact det the. initial cause for the DKA degradatiott is the
excisioE of. pyrimidine dimers frore che DNA !sEelecules vras supported
by the foUowing experimental results that, <1) tbe iinmediate
fragmentation of.. eelZular nNA was ebserved in the eelXs shortly after
the ultravELoiet irradiation by the alkaiine sucrose dradÅ}ent anaiy-
sis, ( 2 ) the pyriwtdine dimers were detected in acid-soluhlLe
Åíractions extractect from the aells inaubated for onZy 5 min after
ultraviolet irradiation, (' 5 ) the post-irradiation degradatioit was'
markedly suppressed in a (ibuble rrvatant,= g. goU.N'G30uvrC, ( 4. )L
eatÅíeine inhibited sigpiÅíicantly the post-irradiation degradation
oÅí ugvA.
The mechapism by whieh the ÅíolLowed extensive DNA degradation'
is resulted from the gaps] in DNA strands. producect by excisiorr of
pyrimidLrie dimera was discussed.
ACKIiTOWLEmm'
( 52 )
rehe author would like to ac.knowiedge the helpful guidance and
critigal diseussions by ProÅíessor Zchijiro Honjo aud Assistsmt Pra.
ressorb Dr. Keiichi Kozu during the preseEt experimeuts.
RIEll!'EREKCES
l. T. Kato and S. Kongo, )futation Res., lt, 2S5 ( 1967 ).
2. I. Kato and S. Kondo,. J. BacterLoL.,. I!,9b &7Z <' L970 ).
5.. I. Kata and S. Kondia. Jap. J. Genet.,Sits. ( Supp. 1 )•, 51 ( 1969 )•
4. A. J. elarls. J. geU. Physiol., ZQg, ( Supp. i ), l65 ( l966 ).
5•. N. S. WUIetts and A. J. erark, J. boeteriol., iOO, 251 ( 1969 ).
6. A. J. Clark anct A.D. Margiutes, Pr'oe. Kati. Aead'. S( t., Z,
451 ( X965 >.
7. P. Eoward-Manders and' L. Theriot., G.enetics, 22, ]L57( 1966 ).
8. ti. Ogawa. K. Shimada arrd J. TeiiKi.zawa. Keiee. Gerr. Genettcs.
,1,,,..Ok 227( 19as )• •
9-. A. J. Clark, M. Chaatberlitt, R. P. Boyce and P. Howard-nanders.
j. )to1. MoZ.. IL2, 442 ( l966 )•.
XO. Z. Horii au-ct K. Suxukt, Photoahem. Photolti;ol., &, 95 (" 1968 ).
(55>
IZ.
12.
15-
-•x5.
16.
17•
1&.
19-
20.
21.
22.
Z. HorLi arrd K. Su2raki, Phptockem. PhotebioX.. ILIt;, 99 ( 1970 >.
I. M.. Eertuetn. Ge"et. Res.. eanb.. Ii,k,,, 291 C 1969 }.
P. Ebward-•F'Xanderg and R. P. Boyee, Racliatiorr Rese SuppZ. J6,
X56 C l966 ).
R. A. McGrath andi R. W. WilLLams. Eature, ,2...12 554 (' 1966 ).
K. C. Siaith. Photoctrem. Photobiol.. J2 50J ( l965 }.
R. B. SetLovt and W. L. carnter, Proc. Natl. Aead. Sei., 21L, 226
( 196it >.
D. S. Ka-pp andi K. C. Smith, J. Bactertei.. iL92, 49 ( 1970 ).
O. haaiLsee artdi P. C. Hanawait. J. Mol. Siol..:5 144 ( l96i )•
K. Shimada ut Y. reaka&i. Bioelutm. HiophJrs. Aeta, llt:E. 765 ( 1967 ).
W. L. Carrier aRd R. B. SetZow, J. ISacteriol.. ,1,,...02.17& ( l970 ).
P. Uoward-nanders. Ami. Rev. ilioekem.. iZ. I75 ( l968 >,.
A. K. Gakesan and K. C. Sntth, J. Bacteriol., 2S6t. 565 ( 1968 ).
C 54 >
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LEGIE gDS FOR F:GURES'
Eig. i. Ultraviolet surutval gurves for !. sigiUii B/r50-R and KG5e
in ditÅíerent cell conditions.
The ultrawtoZet-sensitistity of S. stg,-,li NGI50 in different physiol-
egical conditions were examined. The exauttred cell conditions were
as Åíollows ; growth phase, stationary phase, starved condition Åíor
arnino aeids and a condition pre-incubated at 570C foy 90 !r[i.n with
chlora!nphenieal. g. eoli }l!r50-R was used as control. IÅílr50-R :
stationary, --O- ; growing, - . NG50 : growtng, - ;
statioltary, .--CP ; amino acid-starvation, --"O-' ; incubation with
ahloramphenicoX, - .
Rig. 2. Degradation of the DKA in grewing ceUg of P. sgt,-li NG50
after ultraviolet irradiation. ' I)KA was labelect wLth[H2] -thymidine by incubating the ceils in
]et9 medium suppleittented with 5.5 pC/3 pglml ef[f]-thymadtae and
200 pg/(nt1 deoxyadenosine. The eelZs were wasbect, resuspended in
EM9 medi.um and irradiated with various doses of ultraviolet iight.
The ioss of the radioactivLty from acid-insolubie fractions of the
(i)
cultures duriztg post-irradiation ineubeLtion was detected. The 100
per cent valueg at the zero ineubation time were about 2,OOO cpm/O.2
ml culture. ultraviolet dose :o ergb!rm2, --o- ; 2o ergs/an2,
-o-- ; ioo ergs/ma2, '- ; 4oo ergs/mrn2, - . '
Eig. 5. CE5-thymidine incorporation into acid-insoluble fractions
during incubation in SPS9 medium and M9 rtvediunEe
The cells grown in IN9 medtun were washed, and resuspended in .five fold ve1unn ot M9 mediun. The ceU suspension was divided
into two portions and 2.5 mglal casutno acids were added to the one
portion of the two. tab-thymidine and deoxyadenosine were added
to all the cultures at zero incubation time and the cultures were
incubated at 570C. The experimental details for the measurememt
oÅí ra(lieactivLty incorporated into acid-irtseluale fractions were
described under Material.s etnd Methods. '. ,. i. L' ;..i. ,' .1.'.
""-"-O-- , in EM9 meium ; - , in' M9 medium ; - , celis
were transfered to 1ilM9 medtum after 70 utn starvation-incuhation in
M9 rnedium. '(a> E. coli E!r5Q-R, ( b) E• coli KG50. .te- -
<ii)
1rrg. 4. [di-thyutdine incerporatien into aeid-insolubie Åíractiens
during incubation in us -medium with or without chioranphenicol. ' ' !. coli NG50 was used. The growing cultures in SM9 meditm
vrere supplemented wtth[uJ]-thyntdine and deoxyadenosine,. an-d incu-
bated at j57eC' yeith er yvJ.thout chloranpbenicol. The detatis Åíor
measueement ot[Hl d-thyutdine incorperation were deseribed under
Materials, and Methoct8. - ,wtth chloramphenieol; ---S---
'without chloraiuphenieol.
IeXg. 5. Pest--irradiatiou degradation of' onA in the eelis without
onA replication.
( a ) The asA replieation of S. sigte,sli NG50 was stopped' by agti=o
acid-starvation. The eells which have nc DNA repXication were
ir2adiated ntth wuious doses of ultravLoiet ]i.get. The other 'experimeneai detaiLls were ctescribed unaer Materials and Xetheds.
100 per cent vgiues at zero incubation time were about 1,500 cpm.
-O-- ,non-irradiated; " ,irradia-ted with se ergsZiua2; ' " ,irEadiated witlx loo ergslan2; - ,irradiated witik
2oo ergfitytmm2 ; ---t>' , irradiated with 4oo ergsfma2 ; -s-- .
irradiated with 1,ooe erg.s. /ua2.
'
( "i )
'
( b ) 1!he DNA replication of E. celi Hlr50-R was stopped by
amino acid-starvation and of g.. si2,:ELisi Bsel by chloramphenicol. The
cells vvere irradiated with 1,OOO erg.slmm2. Tbe ZOO per c'eEvt values
at zero incubatio' n- time ver.e about 20oo ut 1200 cpm in ff/r50-R and
Bsel, reSpeetIvely. " ,, Hlr50-R noR-lrr.adiated ; " , 'utr50-R irnadiated with• 1,OOO ergs/mm2 ; -M- , B's-l non-irradiated
; - , Bs-l irradiated vvith- 1.0oo erss!im2.
IMg. 6. Post-irradiation degradation of 1}!ifA in S. sgt,-,li NG50 ceased
the DNA replicatiou by cthloranpheutaol.
EITA replicatien had been stopped through ineubating the cells
'with 50 pg/mi oÅí ehloratnphenicol for 90 ztln. The cultures were
'then irradLated with uarious ctoses of ultraviotet Ught and continued
to incubate at 37ee. The metbods f.or the measurenEent of radio-
actintty in acta"nsoluble frac.tion• were desc eibed under Methoda'.
The iOO per een-t values at zere incubation tinre} were about 1,800
epa. ..'..,.i:"' ---e-- , o erstan2 ; ----e- , src) ergs/rnrn2 ; •--o--"- ,
'loo ergs/izBn2 ; --(>r- , 2oo ergslana ; .--suL- , 4oo ergs/rmi2.
Civ)
M.g. 7. Comparisen of the rate ot pest-irrautation• IMA degradation
in g. sEstl,-,ii NG30 labeled in ditferent osA reeions.
1!we kLnds of Åëells labeied ntth'[R3)-cny!ptdine in difSerertt DNA
region yvithin the onA rrolecules• having no replicating points were
prepared. One is. Zabeled in tt early region ii aiid the other it late
region' ii. The preparation ef the ceZls vere described in detaiZs
under Materials and Hetheds. The cultures were irradiated with
various dopes of ultravi.olet lignt an`d inTcubated at 570C in amino
'acid-free M9 medium. the 100 per cent values at zero ineubation
time were about 90e cprn in the experiments oÅí U early region i} and
1,100 cpm in that ot it iate regton ", respectiveiy. < a ) experi-
ment of N aivly region ", Cb) experiment ot " late region ".
-O-- ,non-irradiated; " ,irradiated with so ergs/mm2;
- ,irradiated utth 100- ergslan2; --7db--,irradiated witit
2oo ergs<fnm2 ; •--t)r-- , irradiated nttn LNpo ergslua2.
IMg. 8. Degradation of MgA in anino acid-starved celis. of g. sEgtaii
KG50s!t!!C afteÅë ultraviolLet irradiation.
S. sigt!iliL EG50!lyt"C vas used. rhe IMA repMcation was stopped
throusb mue acid-starvation. Tbe cells vrere irradiated with
(' v)
sptarious doses of uitravtolet light air.d ineribated at 370C in X9
mediunt. urhe experimental proeedures were the sante as in eig. 5a. 'rke iOO per cent ptues at zere incubatien tine wereAbottt 2}20e elmt.
---o--- ,rron-•ir=autated; " ,i?radiated with so ergofmm2;
--or-- . irractiated with 100 ergs/nrm2L:=2-IN--{ !glradiated witlt 2oO ergsfan2
-; -"'., irradiatect witÅ}t Lioo ergs!mn2.
Iitg. 9. change in the anount eÅí pyrimi(tine diners in the acid-insoluble
fractiorRs of the ceUs of g. gst-,li NG50 azEd NG50uvre during pest-
irraetation imaubatione
S, -tZi NG50 a!td ue3e}ytyne were used. [H;5J-thJrntdine-labeZed
i'ceng were starvea anno acicts, irradiated with 4oo ergs!rma2 and then
ineubated Åíer 60 axkd 120 rnin in X9 nedium. Tlte acia-inseluhlLe
fractiens of the eells w.ere hydrolyzed and pyri!ni(lizte dimers were
deteetea by paperehrouatograplzy. the experimental detaiXs were '
describedi un•der Xetbo• ds. (a>g. sERt,-," ptG3e, (' b) g. s2gttiLli KG30LtC.
-Q--- ,no incultatien; ---e-' .incubated Åíor (so mim ;--.ay-"-',
incubatect for i20 rnin.
C vi. )
Fig. 10. Effect ef cafteine orr DNA degradation in !. eo" NG50
irradiated with ultravioiet light.
MgA replieatien of tke cells was stopped through amino acid'-
starvation. CaÅíÅíeine was added to the euitures and the eells
were irradLated with "Qa ergevtm2 ot ultrautolet light. The
degradation of NA durtsg the post-irradiation incuhation was mea-
sured. The detailg were the same as i!! ptg. Sia except for the
add'iticm of eatteiEe. 100 per e,eitt values at zero incubation
time were about 1.50Q gpm. orat'teine coneentra•sto!t : --tr- .
O per eent ; " , O..i per cent ; r-O--- , 02 per cent ;
---e--- . O.5 per eent.
Vig., U. rniLtial fragmentation of I)KA in the eells' seen after
the ultrautolet irradia•tion.
[llZ5J-thJrmi(tiBe -labeled g. sg.,-.li KG5a and Wr50-R cens were
inauhated at 57eC tor 90 ntll in amit!o acidi-free raediurnt ( M9 medium ).
rhe eeUs weresrradiated with aoo anct 1,ooo ergs/mm2 and ineubated
for 5 mb in M9 "Eedium at 5.7aC. rillmediately after tue incultation.
the cells were coaled, convert.ed into spberoplasts ar!d geEtrtthzged
'in aikaline sucrose gradtene. Tbe. other detaUs were descritsed
( vi'i )
in Materials and Metbocts. C a ) E. celi NG5Q, ( b > g. coli.
Hlr5Q-R. ---(>-- , non-irradrLated ; --e-- , irradiated witli 20Q 'ergmofmm2 ; ------- . iLrradiatea with i..ooo ergs/!ma2.
'
Eig. Z2. fEhe sedirEerttation profiles of DNA of P. ggt-l:L NG30 incubeted
for aO min aft'er uXtrasrcLo Let irradiatu' on.
[LHjll d-thvrrEidine-labeled cells were starvedi aHtuo acidis for 9e mim.
and then irradiated witk various doses of uit-rautolet ligkt.
The irradiate,d aultixves were, incuhated at, 570C Zer 20 utn ta the amino
acid-Åíree medlum. The eells were- lysect bty lysQzyme. iayered on a
allualine sucrose gradient ( 5 - 20 % ). and tben centrifuged- for 90
!gin at 50,OOQ rev.lutn. Neasuremant' ot radioactiutty and the other
'detatls. wgre described under hat•eriais and Methodig. T"o sets: of
experimental results were piled up iE the sane figure.
and ------ , R'on-irpadiatedi ; A--e-` , irradiated with
loo ergermm2 ;•-- --c}- ptb,irradiate•di' with 2oo ergs/mnt2;-.
irradiatedi with 40o ergs/mnt2 ; -"iOr"- , irradiatedi wtth 1,OOO
ergsian2.
( viii )
Fig. Z5. begradation of DNA of S. sEgt-,U NG50 in the course of time
of post--irradta•eian ine.ubatlon-
ltoth stratns, S. sgil,j.i NG50 andi H!r50-R were. usedi. The aells
labelect vvitn[kj)-thymidine' were starvEd uto acids fer 90 mirr anrf
tnen irradiated with 200 or l,OeO ergEtimm2. The irradiated aeUs
weee incubetect at 57eC in' M9 medium Åíore 20 and 120 min. `i'ke ceUs
eonverted into sptieroplasts" were lysed and' layered on tne eop ot
allsaline sucrose gradient. `the otaer details Åíor experiments were
the same as; laefore experiments.
(a)E. Goli NG50, -or ,, non--irradiated; --e--' ,, irradiatect -•, 'witn 2oo ergs/mm2 and incubated fou 2o ntrr ; ---t-- , irradtated
'wivkt 200 ergHs!mra2 anct inauuatect foB 120 utn.
:.'.,.. L. ' vt(b) E. aoli NG350, --O- , Ren-irrtidiatea; --'e-'-',•irradiated . ttwith 1,ooo ergs!mm2 and incubated tor 2o min ;--t-"' ,. irradiated
with 1,ooo ergs/rnm2 and incuDated Åíor i2o min.
( c ) S• eoLi HIV30-R, •---C"--- , non-irradiated ; ---e-- , irradiated
witn 1,ooo ergs/ma2 and incuoated for 2o utn i'-t-- . irradiated
witlt x,poo ergeofma2 and incubatect for 12o min.
(ix)
FiS,. IZg. Dose-dependent. increase in the arnount' of. radioaetivitjr
' Q-Åí acLd-soZubLe fractians aon'taining pyrimidiue diiEers oÅí g.. celi
NG30 ..
[Hjl]-thyrt-idine-labeled eeUs were starvedi for amirEo aeids arrd
thert irrad•tated with various doses. The Åëultures were. inorttbated
Åíor s rm at S7Qe'. Tbe aeict'-soluble fraetions were neutralized'
and' charged to the eolvu!m eÅí 1)owex-1 ( 200 - 400 mesh. X&. er- type ).
After wa$hLng the resin ntth excess volume oÅí water to release fteee
Ere5]-thymndine, 2Q !niL of o.5 K HeL was passed through the eoiuua..
Tlte radioactivity of the eLuates. were caunted.
' Fig. I5. Released pyrimidine diners iEto acia-soiubZe fractions.
S. eoli. NG30 was used. The cell$, starvect amirto acid•s were . i.rradiated with 2,22o ergs!ma2 (' ctgse rate : 74 ergs/rm2/see ), and
then incubated at 57QC Sor S utn. Tbe acid-solikbie fractLo!ts were
eondensed aEd hcydrolyzed with 99 per eent bormie acid at Z7See fo=
X5 pttr. The hydrolysates were then analyxed by paperÅëhremato-
graphy. Emperimental details ara deseribet wnder hateriais and
Xetheds. --<>---- ,, irradiated celXs ; --e-- ,, nen-irradiated cells.
(x)
?Xg. 16. Post-irradi.ation degradation of the IMA in g. eoli NG5a
irE the conditians with or without gLueose.
, rbe eeUs were starved amirra acLds by incubeting tltem Åíor 90 utlt
in M9 mediLum. The irradiated aell& were incubeted' in the M9 medinnk
with or without gZuaose. Tlrve ultraviolet diasfis used were 50 and
loo ergs/mrrL2. r)te otner experimental procedures were the same as
in' Mg. 5a.
- .so ergs/ua2, with giucose;
",so ergs/nn2, without giueose;
--A--- , loo ergslmm2, witlt glueesa ;
+ , loo ergs/umL2,. without glucose.
(' xi- )
TabXe r.
course of
Change in the sedimerrtatiori veeZogity ot
time ot post-irradiation ineubation
DNAmoiecules in
Strains
Dose
( ergs/mm2 )
Post-irradia'-tioE
5
Inc.ubati on (
20'
lstit )
120
ggso
Klr50-R
o
200
looe
o
200
1000
za' ..e
n•5 8'.5
15'.O
i2.0
8.5
12.0
IO .5
7•.5
12 .•S'
-12.0
r 9 ..Q
7'. O
e -•
15.0
The sed'irserrtatierr velocity of DKA
distribution. was: representredl as' a
alkaline suerose grradient•. in' tne
!mplecules- in rmats
distlanc/e frem. the
tigures' ( irr- cm )
peaks of
rsemscus
DNA
oE