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T*
(ancestor)Tj
/F11 1 Tf
3.6286 0 TD
(*)Tj
/F6 1 Tf
1.0022 0 TD
[(1)-289.1(billion)-284.7(years)-289.8(ago,)-290.1(nearly)-291.3(half)-287.9(of)-284.4(essential)-289.9(yeast)]TJ
-4.6308 -1.1174 TD
[(genes)-474.2(are)-466.4(replaceable)-474.3(with)-472.1(the)-471.1(human)-468.7(ortholog)-469.8(\(Kachroo)]TJ
/F7 1 Tf
0 -1.1116 TD
[(et)-481.5(al.)]TJ
/F6 1 Tf
2.2405 0 TD
[(,)-481.5(2015\).)-481.2(Furthe)-7.7(rmore,)]TJ
/F7 1 Tf
9.5438 0 TD
(RAD51)Tj
/F6 1 Tf
3.427 0 TD
[(constitutes)-486.9(part)-483.7(of)-480.2(an)]TJ
-15.2114 -1.1116 TD
[(evolutionarily)-484.9(conserved)-479.7(eukaryotic)-487(pathway)-477.6(for)-481.3(homolo-)]TJ
0 -1.1174 TD
[(gous)-418.9(recombination)-428.3(and)-419.2(recombinational)-426.1(repair)-423.7(\(Baumann)]TJ
0 -1.1116 TD
[(and)-309.7(West,)-316.5(1998\).)-308.4(Due)-310.6(to)-313.3(the)-309.9(translational)-317.7(power)-307(granted)-315.4(by)]TJ
T*
[(this)-367.3(close)-364.9(genetic)-362.8(relationship)-367.2(and)-367.3(the)-361.7(long)-367.4(duration)-361.5(of)-365(the)]TJ
0 -1.1174 TD
[(BioSentinel)-257.7(mission,)-259.9(the)-252.3(DNA)-259.3(damage)-253.2(response)-258.4(observed)-258.6(in)]TJ
0 -1.1116 TD
[(yeast)-336.1(cells)-336.2(will)-334(be)-337.4(directly)-334(relevant)-335.1(to)-336.4(that)-334.1(of)-330.5(astronauts.)]TJ
26.5177 67.9184 TD
[(To)-487.2(measure)-496(the)-488.4(effects)-489.1(of)-491.7(ionizing)-488.4(radiation)-490.7(on)-486(yeast,)]TJ
-1.0137 -1.1174 TD
[(cells)-376.5(are)-374.2(desiccated)-379.8(then)-374.3(rehydrated)-377.4(at)-372.1(specic)-373.8(time)-375.7(points)]TJ
0 -1.1116 TD
[(over)-374.2(the)-367.5(course)-371.6(of)-365(the)-367.5(mission)-371.7(with)-368.4(a)-370.9(mixture)-367.4(of)-365(synthe)-7.7(tic)]TJ
T*
[(complete)-370.9(\(SC\))-368.1(growth)-364.7(medium)-367.5(and)-361.6(alamarBlue)-375.3(\(aB\),)-361.3(a)-365.1(re-)]TJ
T*
[(dox)-412.2(dye)-413.4(that)-409(serves)-417.5(as)-406.3(a)-411.2(metabolism)-419.2(and)-407.6(cell)-415.9(health)-412.4(indi-)]TJ
0 -1.1174 TD
[(cator.)-385.7(Approximately)-387.7(1)]TJ
/F10 1 Tf
9.7627 0 TD
()Tj
/F6 1 Tf
.72 0 TD
(10)Tj
7.3823 0 0 6.5614 426.1038 674.7021 Tm
(5)Tj
9.843 0 0 9.843 433.5306 670.3369 Tm
[(cells)-382.3(are)-385.7(dried)-381.1(inside)-382.2(100-)]TJ
/F11 1 Tf
10.8858 0 TD
(m)Tj
/F6 1 Tf
.5529 0 TD
(L)Tj
-23.6781 -1.1116 TD
[(wells)-266.8(in)-261.5(polycarbonate)-266.9(microuidic)-263.5(cards,)-264.5(two)-263.5(of)-261.4(which)-266.9(will)]TJ
T*
[(be)-268.3(rehydrated)-262.2(at)-262.7(a)-267.2(time.)-263.9(These)-262.3(cards)-266.8(are)-264.8(then)-264.9(assembled)-267.8(into)]TJ
0 -1.1174 TD
[(stacks)-296.7(which)-289.9(include)-292.5(lters,)-293.3(a)-296(thermal)-293.7(control)-291.3(system,)-288.6(and)-292.4(a)]TJ
0 -1.1116 TD
[(three-color)-378.6(LED)-375.2(optical)-379.1(source)-377.3(and)-373.1(detector)-375.4(that)-380.2(takes)-370.6(op-)]TJ
T*
[(tical)-307.7(measurements)-313.8(of)-301.7(the)-309.9(wells.)-304.8(The)-305.2(optical)-309.9(system)-302.5(has)-309.5(the)]TJ
0 -1.1174 TD
[(capability)-400.9(to)-388.2(independently)-398.3(record)-389(changes)-396.9(to)-388.2(turbidity,)-394(as)]TJ
0 -1.1116 TD
[(well)-363.8(as)-366(the)-361.7(colorimetric)-364(changes)-368.1(of)-359.3(aB,)-359.2(enabling)-368.5(both)-361.6(spe-)]TJ
T*
[(cic)-496.3(and)-494(unied)-491.6(interpretat)-10.3(ions)-488.2(of)-497.5(growth)-491.4(and)-494(metabolic)]TJ
T*
[(data.)-315.6(Card)-317.6(stacks)-313.9(are)-316.6(placed)-315.5(on)-307.5(nine-card)-318.7(uidic)-313.1(manifolds,)]TJ
0 -1.1174 TD
[(two)-413.2(of)-388.1(wh)20(ic)19.6(h)-398.5(a)0(r)14(e)-399.7(in)20.7(te)19.6(gr)20.9(at)19.5(ed)-389.3(in)20.7(to)-394(th)20.7(e)-399.7(4)0(U)-389(b)0(i)20.7(o)0(s)21(e)0(n)19.7(s)0(o)15.3(r)-398.5(pa)19.7(yl)20.7(oa)19.7(d)]TJ
0 -1.1116 TD
[(en)13.9(cl)19.5(os)21(ur)20.9(e)-296(a)0(l)13.8(o)0(n)20.8(g)-294.8(wi)19.8(th)-290.3(th)20.7(e)-296(c)0(o)19.7(r)0(r)20.9(e)0(s)19.9(p)0(o)15.1(n)0(d)20.8(i)0(n)20.7(g)-294.8(re)19.7(ag)19.7(en)13.9(t)-295(b)0(a)19.7(g)0(s)21(,)-297.2(tu)20.7(bi)20.7(ng)15.1(,)]TJ
T*
[(pu)15.1(mp)19.6(s,)-516.9(va)19.7(lv)15(es)19.9(,)-527.6(a)0(n)19.7(d)-531(op)20.8(ti)14.8(ca)18.5(l)-525.4(c)0(a)12.8(l)0(i)20.6(b)0(r)20.9(a)0(t)19.5(i)0(o)20.7(n)-531(ce)18.5(ll)20.6(s)-530.8(\()0(F)15.3(i)0(g)20.7(.)-527.6(1A)20(D)20(\).)]TJ
0 -1.1174 TD
[(Attac)-9(hed)-240.6(to)-244.2(the)-240.7(4U)-245(biosensor)-241.2(enclosure,)-247.1(a)-244.2(radiation)-243(sensor)]TJ
0 -1.1116 TD
[(a)-635.8(TimePix-based)-636.4(spectrometerwill)-637.5(measure)-634.3(the)-632.4(linear)]TJ
T*
[(energy)-383.3(transfer)-379.6(\(LET\))-377.7(of)-376.5(the)-373.2(different)-381(ion)-377.8(species)-379.6(that)-380.2(tra-)]TJ
0 -1.1174 TD
[(verse)-335.9(the)-338.7(sensor.)]TJ
1.0137 -1.1116 TD
[(The)-414.6(BioSentinel)-413.3(CubeS)-7.5(at)-412.4(also)-408.7(contains)-414.4(a)-417(2)0(U)-412(spacecraft)]TJ
-1.0137 -1.1116 TD
[(bus,)-294.5(which)-295.7(is)-295.9(solar)-294.6(powered,)-296.5(and)-292.5(houses)-295.4(propulsion,)-291(a)-296(guid-)]TJ
0 -1.1174 TD
[(ance)-301.6(system,)-300.2(data)-300.6(storage,)-301.4(and)-298.2(a)-296(data)-300.6(transponder)-302.3(\(Fig.)-295.8(1E\).)]TJ
0 -1.1116 TD
[(The)-305.2(data)-300.6(gathered)-300.4(from)-305.1(the)-298.3(deep)-300.5(space)-302.5(nanosatellite)-307.4(will)-299.4(be)]TJ
T*
[(telemetered)-221.1(back)-219.8(to)-215.4(Earth)-215.3(and)-217.6(compared)-215.1(to)-215.4(both)-217.6(synchronous)]TJ
T*
[(ground)-329.1(controls)-332.6(and)-327(the)-332.9(ISS)-325.5(payload,)-330.4(which)-330.2(will)-334(serve)-330.2(as)-325.7(a)]TJ
0 -1.1174 TD
[(microgravity)-351.2(control)-348.9(in)-342.1(addition)-350.2(to)-342.2(a)-347.8(probe)-346.5(for)-343.1(this)-344.3(distinct)]TJ
0 -1.1116 TD
[(radiation)-346.7(environment.)-347.6(For)-342.9(each)-342(payload,)-347.7(yeast)-341.8(growth)-347.4(and)]TJ
T*
[(metabolism)-200.3(rates)-192(will)-195.7(be)-193.4(correlated)-200.2(with)-189.8(the)-194.7(onboard)-193.2(physical)]TJ
0 -1.1174 TD
[(radiation)-294.8(dosimetry)-294.6(and)-292.4(spectroscopy.)]TJ
1.0137 -1.1116 TD
[(Th)19.6(e)-255.7(b)0(i)20.7(g)0(g)15.1(e)0(s)19.9(t)-254.7(sc)19.9(ie)19.5(nt)20.7(i)15.1(c)-255.7(r)0(i)20.7(s)0(k)-243.9(t)0(o)-244.2(t)0(h)20.7(e)-255.7(su)15.3(cc)18.5(es)19.9(s)-254.3(o)0(f)-244.1(t)0(h)20.7(e)-255.7(Bi)19.7(oS)15.2(en)19.7(ti)20.6(ne)19.7(l)]TJ
-1.0137 -1.1116 TD
[(mi)13.7(ss)21.2(io)20.7(n)-312.1(i)0(s)-301.6(l)0(o)15(n)0(g)20.8(-)0(t)20.7(e)0(r)19.7(m)-313.3(ce)18.5(ll)-302(su)15.3(rv)20.9(iv)20.7(al)19.5(.)-314.5(C)0(o)19.8(m)0(p)19.6(a)0(r)19.7(e)0(d)-302.9(t)0(o)-307.6(h)0(u)20.8(m)0(a)18.4(n)-312.1(ce)18.5(ll)20.6(s)]TJ
0 -1.1174 TD
[(an)13.9(d)-335.2(m)0(a)18.4(n)0(y)-330.5(o)0(t)20.7(h)0(e)19.7(r)-340.9(mo)19.6(de)19.7(l)-335.3(o)0(r)20.9(g)0(a)13.9(n)0(i)20.7(s)0(m)19.8(s)0(,)-326.9(y)0(e)19.7(a)0(s)14.1(t)-335.3(is)-330.4(id)20.7(ea)18.5(ll)20.6(y)-340.9(s)0(u)21(i)0(t)20.6(e)0(d)-331.7(f)0(o)20.9(r)]TJ
0 -1.1116 TD
[(pr)15.1(ol)20.7(on)20.8(ge)19.7(d)-237.2(b)0(i)20.7(o)0(s)21(e)0(n)13.9(s)0(o)21(r)-237.2(mi)19.5(ss)21.2(io)20.7(n)-237.2(d)0(u)20.8(r)0(a)14(t)0(i)20.6(o)0(n)20.8(s)-237(du)20.8(e)-238.4(t)0(o)-227(i)0(t)20.6(s)-237.1(ab)19.7(il)20.6(it)20.6(y)-237.2(t)0(o)-227(u)0(s)21(e)]TJ
T*
[(an)13.9(hy)20.8(dr)20.9(ob)20.8(io)20.7(si)15.1(s.)-303.8(T)15(h)0(i)14.9(s)-311.9(ra)19.7(re)-302.8(an)19.7(d)-312.1(r)0(e)14(m)15(ar)14(ka)19.7(bl)20.7(e)-313.3(s)0(t)20.9(r)0(a)19.7(t)0(e)13.8(g)0(y)20.8(,)-308.7(al)13.8(so)-301.5(fo)20.9(un)20.8(d)]TJ
T*
[(in)-526.5(or)15.1(ga)19.7(ni)20.7(sm)19.8(s)-536.6(s)0(u)21(c)0(h)-527.5(a)0(s)-527.3(t)0(a)19.5(r)0(d)20.9(i)0(g)20.7(r)0(a)14(d)0(e)19.7(s)-536.6(\(B)19.8(oo)20.8(th)20.7(by)]TJ
/F7 1 Tf
18.719 0 TD
[(et)-527.6(al)20.7(.)]TJ
/F6 1 Tf
2.2578 0 TD
-.0208 Tc
[(,)-554.1(2)-20.8(01)-20.8(7\))-20.8(,)]TJ
-20.9769 -1.1174 TD
0 Tc
[(ne)13.9(ma)18.4(to)20.7(de)19.7(s)-288.9(\()-70.2(Ma)18.8(di)14.9(n)-283.3(a)0(n)13.9(d)-283.3(Cr)14.1(ow)20(e,)-276.4(19)20.8(75)20.8(\),)-281(in)20.7(se)19.9(ct)13.8(s)-283.1(\()0(S)15.3(a)0(k)19.7(u)0(r)20.9(a)0(i)]TJ
/F7 1 Tf
22.0366 0 TD
[(et)-280(al)20.7(.)]TJ
/F6 1 Tf
2.0101 0 TD
(,)Tj
-24.0468 -1.1116 TD
[(20)15.1(08)20.8(\),)-459.5(an)19.7(d)-467.6(p)0(r)20.9(o)0(k)20.8(a)0(r)19.7(y)0(o)20.8(t)0(e)13.8(s)-467.4(\(P)21(ot)20.7(ts)20.9(,)-470(1)0(9)20.8(9)0(4)20.8(\))0(,)-459.5(a)0(l)19.5(l)0(o)20.7(w)0(s)-457.9(o)0(r)15.1(g)0(a)19.7(n)0(i)20.7(s)0(m)19.8(s)-467.4(to)]TJ
T*
[(su)15.3(rv)20.9(iv)20.7(e)-514.9(e)0(x)19.7(t)0(r)15(e)0(m)18.5(e)-509.1(de)13.9(si)20.9(cc)18.5(at)19.6(io)20.7(n)-513.7(a)0(n)19.7(d)-513.7(re)14(ve)19.7(rs)21.1(ib)20.7(ly)-503.4(ha)13.9(lt)-503.5(me)18.4(ta)19.6(bo)20.8(li)20.6(c)]TJ
0 -1.1174 TD
[(ac)12.8(ti)20.6(vi)20.7(ty)-313.3(un)20.8(de)19.7(r)-323.6(t)0(h)20.7(e)0(s)14.1(e)-324.8(co)19.7(nd)20.8(it)20.6(io)20.7(ns)-313(\(D)14.2(up)20.8(on)20.8(t)]TJ
/F7 1 Tf
16.1963 0 TD
[(et)-314.5(al)20.7(.)]TJ
/F6 1 Tf
2.0447 0 TD
.3202 Tc
[(,2)320.2(0)341(1)320.2(4)335.3(\))320.2(.)]TJ
-17.2273 -1.1116 TD
.0117 Tc
[(Du)4.8(e)-405.3(t)-1(o)-398.3(i)-1(t)4.7(s)-403.9(co)4.8(lo)4.8(nizat)4.7(i)-1(o)4.8(n)-404.1(of)-398.3(terrest)4.7(rial)-404.2(i)4.7(n)-.9(terf)4.9(acial)-404.2(n)-.9(i)4.7(c)-2.1(h)4.8(e)-2.1(s,)]TJ
-1.0137 -1.1116 TD
.0122 Tc
[(inclu)5.3(d)-.4(i)5.2(n)-.4(g)-432.4(t)5.2(he)-433.6(s)5.5(u)-.4(r)5.4(f)-.4(ace)-433.6(o)5.3(f)-432.4(s)-.2(o)5.3(il,)-429(r)5.4(o)-.4(ck)5.3(s,)-429(pl)5.2(an)5.3(ts,)-429(a)4.2(nd)-432.4(ani)5.2(m)-1.6(als,)]TJ
/F7 1 Tf
0 -1.1174 TD
.01 Tc
[(S.)-460(ce)-3.8(re)-3.8(visiae)]TJ
/F6 1 Tf
5.8634 0 TD
.0117 Tc
[(i)4.7(s)-455.7(oft)4.7(en)-455.9(s)5(ubj)4.7(ected)-455.9(t)-1(o)-455.9(s)-.7(evere)-457.1()5(uct)4.7(u)-.9(ati)4.7(o)-.9(n)4.8(s)-461.5(i)4.7(n)]TJ
-5.8634 -1.1116 TD
.0119 Tc
[(hyd)5(rati)4.9(on.)-458.1(As)-461.3(y)5(eas)5.2(t)-461.6(c)-1.9(ell)4.9(s)-467.1(u)5(n)-.7(d)5(e)-1.9(r)5.1(g)-.7(o)-461.5(d)-.7(esi)4.9(c)-1.9(cat)4.9(i)-.9(on)5(,)-463.8(t)4.9(hey)-467.3(f)5.1(ace)]TJ
T*
[(mech)5.1(ani)5(c)-1.8(al,)-279.4(s)-.4(t)5(r)-.6(u)5.1(c)-1.8(t)5(ural,)-279.4(an)5.1(d)-282.8(o)5.1(xi)5(dat)5(i)-.7(v)5.1(e)-284(challeng)5.1(es)5.3(,)-285.2(i)5(nclu)5.1(din)5.1(g)]TJ
0 -1.1174 TD
.0117 Tc
[(int)4.7(r)-.9(acellu)4.8(lar)-490.5(c)3.7(rowdi)4.7(ng)4.8(,)-492.8(p)4.8(la)]TJ
11.3581 0 TD
.0113 Tc
[(sma)-492.1(m)-2.5(embrane)-486.3(lys)4.6(i)-1.4(s)-490.7(a)3.3(nd)-485.1(per-)]TJ
-11.3581 -1.1116 TD
.0125 Tc
[(meab)5.6(ili)5.5(zati)5.5(on)5.6(,)-336.5(s)5.8(tru)5.6(c)-1.3(t)5.5(u)-.1(r)5.7(a)-1.3(l)-334.3(m)4.4(od)5.6(i)5.8(catio)5.6(ns)-334(t)5.5(o)-334.2(p)5.6(rot)5.5(ei)5.5(ns)5.8(,)-336.5(p)5.6(ro)5.6(tei)5.5(n)]TJ
T*
.012 Tc
[(agg)5.1(r)-.6(egati)5(on,)-198.8(lip)5.1(id)-202.2(p)5.1(h)-.6(ase)-203.4(t)-.7(r)5.2(a)-1.8(ns)5.3(it)5(io)5.1(ns,)-198.8(a)-1.8(nd)-202.2(o)5.1(x)-.6(i)5(d)-.6(ativ)5.1(e)-203.4(s)-.4(tr)5.2(es)5.3(s.)-204.5(To)]TJ
T*
.007 Tc
[(s)-5.4(u)-5.6(rv)-5.6(iv)-5.6(e)-450.3(t)-5.7(he)-6.8(se)-450.3(i)-5.7(n).1(s)-5.4(u)-5.6(lt)-5.7(s,)-451.4(y)-5.6(e)-1(a)-6.8(s).3(t)-449.2(h)-5.6(a)-6.8(s)-448.9(ev)-5.6(ol)-5.7(ve)-6.8(d)-449.1(a)-450.3(v)-5.6(a)-1(r)-5.6(i)0(e)-6.8(t)0(y)-449.1(o)-5.6(f)-449.1(e)-6.8(n-)]TJ
0 -1.1174 TD
.0127 Tc
[(dog)5.8(en)5.8(ou)5.8(s)-604.5(p)5.8(ro)5.8(tecti)5.7(v).1(e)-600.1(m)4.6(echan)5.8(i)0(s)6(m)-1.1(s)6(,)-601.3(in)5.8(cl)5.7(udi)5.7(ng)-598.9(a)-600.1(u).1(n)5.8(i)0(qu)5.8(el)5.7(y)]TJ
0 -1.1116 TD
.0117 Tc
[(elast)4.7(i)-1(c)-301.6(c)3.7(el)4.7(l)-300.6(w)-1.8(all,)-297(th)4.8(e)-301.6(a)-2.1(ccu)4.8(m)-2.1(u)4.8(l)-1(ati)4.7(o)-.9(n)-300.4(o)-.9(f)-300.4(i)4.7(nt)4.7(racellu)4.8(lar)-300.4(g)-.9(l)4.7(y)-.9(cero)4.8(l)]TJ
T*
.012 Tc
[(and)-288.6(t)-.7(r)5.2(e)-1.8(halo)5.1(se,)-296.7(a)4(nd)-294.4(t)5(h)-.6(e)-289.8(i)-.7(n)5.1(ducti)5(on)-294.4(o)5.1(f)-294.3(st)5(ress)-294.2(p)5.1(rot)5(ei)5(ns)-294.2(and)-288.6(b)-.6(io)5.1(-)]TJ
0 -1.1174 TD
.0095 Tc
[(c)-4.3(h)-3.1(emic)-4.3(al)-216.4(antioxida)-4.3(t)2.5(ive)-217.4(s)-2.9(yste)]TJ
11.577 0 TD
.2246 Tc
[(m)216.5(s)212.2(.R)211(e)216.6(h)212(y)217.7(d)212(r)217.8(a)210.8(t)217.6(i)211.9(o)212(n)4.6(c)210.8(a)216.6(na)216.6(l)211.9(s)217.9(oc)216.6(a)210.8(u)212(s)217.9(e)]TJ
-11.577 -1.1116 TD
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T*
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T*
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T*
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ET
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BT
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[(FIG.)-244.6(1.)]TJ
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