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Sergei Y. Funikov, Dina A. Kulikova, George S. Krasnov, Alexander P. Rezvykh, Lubov N. Chuvakova, Natalia G. Shostak, Elena S. Zelentsova, Justin P. Blumenstiel, Michael B. Evgen’ev
Spontaneous gain of susceptibility suggests a novel mechanism of resistance to hybrid dysgenesis in Drosophila virilis
www.plosgenetics.org
www.plosgenetics.org
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280.8 537.3354 l
h
f*
283.0677 537.3354 m
287.8299 537.3354 l
h
f*
290.0976 537.3354 m
294.8598 537.3354 l
h
f*
297.0709 537.3354 m
306.5953 537.3354 l
h
f*
200.0126 328.9323 m
292.1386 328.9323 l
h
f*
465.8457 263.9055 m
485.4047 263.9055 l
h
f*
515.9055 250.9228 m
525.3732 250.9228 l
h
f*
527.6409 250.9228 m
537.1654 250.9228 l
h
f*
539.4331 250.9228 m
548.9008 250.9228 l
h
f*
252.9071 211.9181 m
262.4315 211.9181 l
h
f*
431.263 185.8961 m
440.7874 185.8961 l
h
f*
443.0551 185.8961 m
452.5795 185.8961 l
h
f*
454.8472 185.8961 m
464.315 185.8961 l
h
f*
466.5827 185.8961 m
476.1071 185.8961 l
h
f*
478.3748 185.8961 m
487.8425 185.8961 l
h
f*
379.8992 146.948 m
389.4236 146.948 l
h
f*
463.0677 120.926 m
474.6898 120.926 l
h
f*
479.1118 120.926 m
489.6567 120.926 l
h
f*
494.1354 120.926 m
545.2724 120.926 l
h
f*
0 g
1 j
1 J
0 w
10 0 0 10 200.0125 707.4141 cm
BT
/F7 1 Tf
1 TL
-0.004 Tc
0 0 Td
(target)Tj
2.5114 0 Td
(specific)Tj
3.1862 0 Td
(inducing)Tj
3.7984 0 Td
(TE)Tj
1.4003 0 Td
(familiesapplies)Tj
7.0185 0 Td
(in)Tj
/F8 1 Tf
1.0318 0 Td
[()]TJ
/F7 1 Tf
(.)Tj
/F8 1 Tf
1.1395 0 Td
[()-13()12()-9()4()12()-4()]TJ
/F7 1 Tf
2.3868 0 Td
([)Tj
0.83 0.64 0.02 0 k
(3)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(4)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(18)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(19)Tj
0 g
(].)Tj
4.6544 0 Td
(This)Tj
1.9843 0 Td
(is)Tj
0.8504 0 Td
(because)Tj
3.3165 0 Td
(several)Tj
-33.2785 -1.3039 Td
(neutral)Tj
3.855 0 Td
(\()Tj
/F8 1 Tf
[()]TJ
/F7 1 Tf
(\))Tj
1.5931 0 Td
(strains)Tj
2.863 0 Td
(exhibit)Tj
2.9763 0 Td
(immunity)Tj
5.006 0 Td
(to)Tj
1.0261 0 Td
(HD)Tj
1.7235 0 Td
(in)Tj
1.0318 0 Td
(dysgenic)Tj
3.6907 0 Td
(crosses)Tj
3.033 0 Td
(but)Tj
1.5534 0 Td
(lack)Tj
1.8198 0 Td
(maternal)Tj
3.7928 0 Td
(piRNA)Tj
-33.9645 -1.2982 Td
(corresponding)Tj
6.0604 0 Td
(to)Tj
/F8 1 Tf
1.0205 0 Td
[()23(\r)2()6(\r)15()4()11()4(\r)]TJ
/F7 1 Tf
3.651 0 Td
(elements,)Tj
3.9911 0 Td
(the)Tj
1.4683 0 Td
(presumptive)Tj
5.2044 0 Td
(primary)Tj
3.4526 0 Td
(driver)Tj
2.6362 0 Td
(of)Tj
1.0148 0 Td
(dysgenesis)Tj
4.4107 0 Td
([)Tj
0.83 0.64 0.02 0 k
(19)Tj
0 g
(].)Tj
2.0806 0 Td
(If)Tj
/F8 1 Tf
-34.9906 -1.2983 Td
[()23(\r)2()6(\r)15()4()11()4(\r)]TJ
/F7 1 Tf
3.6453 0 Td
(is)Tj
0.8447 0 Td
(a)Tj
0.652 0 Td
(key)Tj
1.593 0 Td
(driver)Tj
2.6362 0 Td
(of)Tj
1.0148 0 Td
(dysgenesis,)Tj
4.6375 0 Td
(how)Tj
1.9332 0 Td
(do)Tj
1.2529 0 Td
(neutral)Tj
3.0671 0 Td
(strains)Tj
2.8629 0 Td
(exhibit)Tj
2.9707 0 Td
("immunity")Tj
4.8643 0 Td
(in)Tj
1.0318 0 Td
(the)Tj
1.4683 0 Td
(absence)Tj
-34.4747 -1.2983 Td
(of)Tj
1.0147 0 Td
(maternally)Tj
4.4958 0 Td
(transmitted)Tj
/F8 1 Tf
4.8699 0 Td
[()23(\r)2()6(\r)15()4()11()4(\r)]TJ
/F7 1 Tf
3.6453 0 Td
(piRNA?)Tj
3.4185 0 Td
(Two)Tj
2.024 0 Td
(fundamental)Tj
5.3177 0 Td
(issues)Tj
2.5172 0 Td
(arise.)Tj
2.3017 0 Td
(First,)Tj
2.2734 0 Td
(as)Tj
1.0148 0 Td
(observed)Tj
3.7927 0 Td
(in)Tj
/F8 1 Tf
-36.6857 -1.3039 Td
[()]TJ
/F7 1 Tf
(.)Tj
/F8 1 Tf
1.1395 0 Td
[()6(\r)15()4( )16()6()2()2( )9()11(\r)2()]TJ
/F7 1 Tf
(,)Tj
5.6125 0 Td
(is)Tj
0.8448 0 Td
(there)Tj
2.2677 0 Td
(a)Tj
0.6519 0 Td
(single)Tj
2.5285 0 Td
(major)Tj
2.6135 0 Td
(element)Tj
3.4016 0 Td
(that)Tj
1.7801 0 Td
(serves)Tj
2.6136 0 Td
(as)Tj
1.0148 0 Td
(a)Tj
0.6519 0 Td
(key)Tj
1.5931 0 Td
(driver)Tj
2.6362 0 Td
(of)Tj
1.0148 0 Td
(HD)Tj
1.7234 0 Td
(in)Tj
/F8 1 Tf
1.0375 0 Td
[()]TJ
/F7 1 Tf
(.)Tj
/F8 1 Tf
1.1395 0 Td
[()-13()12()-9()4()12()-4()]TJ
/F7 1 Tf
(?)Tj
-34.2649 -1.2982 Td
(Second,)Tj
3.3505 0 Td
(do)Tj
/F8 1 Tf
1.2529 0 Td
[()]TJ
/F7 1 Tf
(strains)Tj
3.8891 0 Td
(confer)Tj
2.7779 0 Td
(their)Tj
2.109 0 Td
(resistance)Tj
4.1329 0 Td
(to)Tj
1.0261 0 Td
(HD)Tj
1.7235 0 Td
(solely)Tj
2.4548 0 Td
(through)Tj
3.4469 0 Td
(maternally)Tj
4.4957 0 Td
(provisioned)Tj
-30.6593 -1.2983 Td
(piRNA)Tj
3.0443 0 Td
(or)Tj
1.0999 0 Td
(through)Tj
3.4469 0 Td
(alternate)Tj
3.6963 0 Td
(mechanisms?)Tj
5.5729 0 Td
(Despite)Tj
3.2485 0 Td
(significant)Tj
4.388 0 Td
(progress)Tj
3.6 0 Td
(in)Tj
1.0375 0 Td
(understanding)Tj
6.0661 0 Td
(the)Tj
-35.2004 -1.3039 Td
(morphogenetic)Tj
6.3155 0 Td
(events)Tj
2.7326 0 Td
(occurring)Tj
4.1272 0 Td
(during)Tj
2.9253 0 Td
(gametogenesis)Tj
6.0151 0 Td
(and)Tj
1.7348 0 Td
(embryogenesis)Tj
6.1512 0 Td
(in)Tj
1.0374 0 Td
(the)Tj
1.4684 0 Td
(progeny)Tj
3.5092 0 Td
(of)Tj
/F8 1 Tf
-36.0167 -1.2983 Td
[()]TJ
/F7 1 Tf
(.)Tj
/F8 1 Tf
1.1395 0 Td
[()-13()12()-9()4()12()-4()]TJ
/F7 1 Tf
2.3811 0 Td
(dysgenic)Tj
3.6906 0 Td
(crosses,)Tj
3.2542 0 Td
(these)Tj
2.2564 0 Td
(questions)Tj
4.0195 0 Td
(still)Tj
1.6384 0 Td
(need)Tj
2.143 0 Td
(to)Tj
1.0261 0 Td
(be)Tj
1.1508 0 Td
(answered)Tj
3.9912 0 Td
([)Tj
0.83 0.64 0.02 0 k
(11)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(18)Tj
0 g
(].)Tj
-25.4946 -1.2983 Td
(To)Tj
1.3379 0 Td
(answer)Tj
3.0444 0 Td
(these)Tj
2.2564 0 Td
(questions,)Tj
4.2462 0 Td
(by)Tj
1.1792 0 Td
(using)Tj
2.3925 0 Td
(small)Tj
2.3244 0 Td
(RNA)Tj
2.262 0 Td
(deep-sequencing)Tj
6.9562 0 Td
(and)Tj
1.7348 0 Td
(qPCR,)Tj
2.7949 0 Td
(we)Tj
1.3266 0 Td
(decided)Tj
3.3392 0 Td
(to)Tj
-36.3909 -1.3039 Td
(perform)Tj
3.5149 0 Td
(a)Tj
0.6576 0 Td
(comparative)Tj
5.1704 0 Td
(survey)Tj
2.8119 0 Td
(of)Tj
1.0148 0 Td
(maternal)Tj
3.7928 0 Td
(piRNA)Tj
3.05 0 Td
(profiles)Tj
3.1975 0 Td
(across)Tj
2.6815 0 Td
(several)Tj
2.931 0 Td
(neutral)Tj
3.8551 0 Td
(strains)Tj
2.863 0 Td
(of)Tj
-35.5405 -1.2982 Td
(different)Tj
3.6566 0 Td
(origin)Tj
2.6419 0 Td
(that)Tj
1.7801 0 Td
(did)Tj
1.5421 0 Td
(not)Tj
1.576 0 Td
(quite)Tj
2.2451 0 Td
(fit)Tj
1.0714 0 Td
(the)Tj
1.4684 0 Td
(HD)Tj
1.7234 0 Td
(paradigm)Tj
4.0592 0 Td
(developed)Tj
4.269 0 Td
(in)Tj
1.0374 0 Td
(the)Tj
1.4684 0 Td
(previous)Tj
3.651 0 Td
(studies)Tj
2.982 0 Td
(of)Tj
-35.172 -1.2983 Td
(this)Tj
1.6724 0 Td
(phenomenon)Tj
5.5899 0 Td
([)Tj
0.83 0.64 0.02 0 k
(3)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(4)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(9)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
(19)Tj
0 g
(].)Tj
4.1839 0 Td
(Additionally,)Tj
5.4538 0 Td
(we)Tj
1.3266 0 Td
(developed)Tj
4.269 0 Td
(transgenic)Tj
4.3653 0 Td
(strains)Tj
2.8573 0 Td
(containing)Tj
4.5297 0 Td
(a)Tj
0.652 0 Td
(pre-)Tj
-34.8999 -1.3039 Td
(sumptive)Tj
3.8891 0 Td
(causative)Tj
3.8437 0 Td
(TE)Tj
1.4003 0 Td
(and)Tj
1.7291 0 Td
(did)Tj
1.5421 0 Td
(not)Tj
1.576 0 Td
(detect)Tj
2.6136 0 Td
(a)Tj
0.6519 0 Td
(cytotype)Tj
3.6 0 Td
(change)Tj
3.0444 0 Td
(after)Tj
2.0353 0 Td
(its)Tj
1.1451 0 Td
(propagation)Tj
5.0854 0 Td
(in)Tj
1.0374 0 Td
(the)Tj
-33.1934 -1.2983 Td
(genome.)Tj
3.6339 0 Td
(The)Tj
1.7802 0 Td
(accumulated)Tj
5.3064 0 Td
(data)Tj
1.9106 0 Td
(failed)Tj
2.4094 0 Td
(to)Tj
1.0261 0 Td
(pinpoint)Tj
3.6907 0 Td
(a)Tj
0.652 0 Td
(single)Tj
2.5341 0 Td
(TE)Tj
1.4003 0 Td
(or)Tj
1.0999 0 Td
(specific)Tj
3.1861 0 Td
(set)Tj
1.3039 0 Td
(of)Tj
1.0148 0 Td
(TEs)Tj
1.7632 0 Td
(responsible)Tj
-32.7116 -1.2982 Td
(for)Tj
1.3832 0 Td
(their)Tj
2.109 0 Td
(immunity)Tj
5.006 0 Td
(and)Tj
1.7348 0 Td
(support)Tj
3.3165 0 Td
(a)Tj
0.6576 0 Td
(model)Tj
2.7439 0 Td
(in)Tj
1.0318 0 Td
(which)Tj
2.6532 0 Td
(resistance)Tj
4.1329 0 Td
(to)Tj
1.0205 0 Td
(TE-mediated)Tj
5.4765 0 Td
(sterility)Tj
3.2031 0 Td
(during)Tj
-34.469 -1.2983 Td
(dysgenesis)Tj
4.4106 0 Td
(may)Tj
1.9276 0 Td
(be)Tj
1.1508 0 Td
(achieved)Tj
3.7021 0 Td
(by)Tj
1.1848 0 Td
(a)Tj
0.652 0 Td
(mechanism)Tj
4.8472 0 Td
(that)Tj
1.7745 0 Td
(varies)Tj
2.5341 0 Td
(across)Tj
2.6759 0 Td
(strains.)Tj
3.0898 0 Td
(We)Tj
1.6044 0 Td
(thus)Tj
1.9389 0 Td
(propose)Tj
3.4242 0 Td
(an)Tj
-34.9169 -1.3039 Td
(alternate)Tj
3.6963 0 Td
(model)Tj
2.7439 0 Td
(to)Tj
1.0262 0 Td
(explain)Tj
3.1237 0 Td
(resistance)Tj
4.1272 0 Td
(to)Tj
1.0262 0 Td
(TE)Tj
1.4003 0 Td
(mediated)Tj
3.9458 0 Td
(sterility)Tj
3.2031 0 Td
(in)Tj
/F8 1 Tf
1.0375 0 Td
[()]TJ
/F7 1 Tf
(.)Tj
/F8 1 Tf
1.1395 0 Td
[()-13()12()-9()4()12()-4()]TJ
/F7 1 Tf
(.)Tj
2.6079 0 Td
(Instead)Tj
3.1521 0 Td
(of)Tj
1.0148 0 Td
(solely)Tj
-33.2445 -1.2983 Td
(being)Tj
2.4321 0 Td
(explained)Tj
4.0705 0 Td
(by)Tj
1.1849 0 Td
(maternal)Tj
3.7927 0 Td
(piRNAs)Tj
3.4072 0 Td
(that)Tj
1.7802 0 Td
(target)Tj
2.5114 0 Td
(inducing)Tj
3.7928 0 Td
(TE)Tj
1.4003 0 Td
(families,)Tj
3.5546 0 Td
(the)Tj
1.4683 0 Td
(chromatin)Tj
4.4164 0 Td
(profile)Tj
2.8403 0 Td
(of)Tj
-36.6517 -1.2983 Td
(repeats)Tj
3.05 0 Td
(in)Tj
1.0318 0 Td
(the)Tj
1.4684 0 Td
(maternal)Tj
3.7984 0 Td
(genome)Tj
3.4072 0 Td
(may)Tj
1.9275 0 Td
(confer)Tj
2.7836 0 Td
(general)Tj
3.1295 0 Td
(immunity)Tj
4.2236 0 Td
(to)Tj
1.0261 0 Td
(the)Tj
1.4683 0 Td
(harmful)Tj
3.4356 0 Td
(effects)Tj
2.7269 0 Td
(of)Tj
1.0148 0 Td
(TE)Tj
-34.4917 -1.3039 Td
(mobilization.)Tj
ET
Q
q
1 j
1 J
0 w
11.9999 0 0 11.9999 200.0125 405.0141 cm
BT
/F3 1 Tf
1 TL
-0.0033 Tc
0 0 Td
(Results)Tj
3.7984 0 Td
(and)Tj
1.989 0 Td
(discussion)Tj
/F14 1 Tf
-5.7874 -1.5023 Td
(A)Tj
0.9212 0 Td
(survey)Tj
2.9197 0 Td
(of)Tj
1.0678 0 Td
(neutral)Tj
3.2456 0 Td
(strain)Tj
2.6552 0 Td
(piRNA)Tj
3.1465 0 Td
(profiles)Tj
3.4016 0 Td
(reveals)Tj
3.0803 0 Td
(that)Tj
1.9134 0 Td
(there)Tj
2.3764 0 Td
(is)Tj
0.8882 0 Td
(no)Tj
1.2992 0 Td
(common)Tj
-26.9151 -1.167 Td
(pattern)Tj
3.3071 0 Td
(of)Tj
1.0677 0 Td
(maternal)Tj
4.0016 0 Td
(piRNAs)Tj
3.5292 0 Td
(derived)Tj
3.3401 0 Td
(from)Tj
2.3103 0 Td
(asymmetric)Tj
5.0977 0 Td
(TEs)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 200.0125 355.9748 cm
BT
/F7 1 Tf
1 TL
-0.004 Tc
0 0 Td
(To)Tj
1.3379 0 Td
(characterize)Tj
5.0286 0 Td
(the)Tj
1.4684 0 Td
(piRNA)Tj
3.05 0 Td
(profiles)Tj
3.1975 0 Td
(across)Tj
2.6815 0 Td
(diverse)Tj
3.0501 0 Td
(strains)Tj
2.8573 0 Td
(that)Tj
1.7801 0 Td
(vary)Tj
1.9333 0 Td
(in)Tj
1.0374 0 Td
(resistance)Tj
4.1329 0 Td
(to)Tj
1.0205 0 Td
(HD,)Tj
1.9502 0 Td
(we)Tj
1.3266 0 Td
(per-)Tj
-35.8523 -1.2982 Td
(formed)Tj
3.1577 0 Td
(small)Tj
2.3244 0 Td
(RNA)Tj
2.2621 0 Td
(sequencing)Tj
4.7168 0 Td
(on)Tj
1.2756 0 Td
(six)Tj
/F8 1 Tf
1.3152 0 Td
[()]TJ
/F7 1 Tf
(.)Tj
/F8 1 Tf
1.1395 0 Td
[()-13()12()-9()4()12()-4()]TJ
/F7 1 Tf
2.3811 0 Td
(strains)Tj
2.8573 0 Td
(obtained)Tj
3.7361 0 Td
(from)Tj
2.2053 0 Td
(various)Tj
3.1408 0 Td
(sources)Tj
3.1974 0 Td
(\(see)Tj
0.83 0.64 0.02 0 k
-33.7093 -1.2983 Td
(Materials)Tj
3.9458 0 Td
(and)Tj
1.7291 0 Td
(Methods)Tj
0 g
(\))Tj
4.0988 0 Td
(and)Tj
1.7292 0 Td
(maintained)Tj
4.7905 0 Td
(in)Tj
1.0375 0 Td
(our)Tj
1.627 0 Td
(laboratory)Tj
4.3484 0 Td
(for)Tj
1.3833 0 Td
(more)Tj
2.3414 0 Td
(than)Tj
2.0296 0 Td
(20)Tj
1.1735 0 Td
(years.)Tj
2.4888 0 Td
(These)Tj
-32.7229 -1.2982 Td
(strains)Tj
2.8573 0 Td
(exhibit)Tj
2.9763 0 Td
(different)Tj
3.6567 0 Td
(levels)Tj
2.3754 0 Td
(of)Tj
1.0148 0 Td
(gonadal)Tj
3.3845 0 Td
(atrophy)Tj
3.3279 0 Td
(when)Tj
2.4038 0 Td
(crossed)Tj
3.1974 0 Td
(with)Tj
1.9956 0 Td
(males)Tj
2.5058 0 Td
(of)Tj
/F8 1 Tf
1.0148 0 Td
[()]TJ
/F7 1 Tf
(-like)Tj
2.5512 0 Td
(strain)Tj
/F8 1 Tf
2.5001 0 Td
[()]TJ
/F7 1 Tf
(.)Tj
-35.7616 -1.304 Td
(Two)Tj
2.0182 0 Td
(of)Tj
1.0148 0 Td
(them)Tj
2.2904 0 Td
(\()Tj
/F8 1 Tf
[()]TJ
/F7 1 Tf
1.0261 0 Td
(and)Tj
/F8 1 Tf
1.7291 0 Td
[()]TJ
/F7 1 Tf
(\))Tj
1.4911 0 Td
(represent)Tj
3.9571 0 Td
(strong)Tj
/F8 1 Tf
2.7723 0 Td
[()]TJ
/F7 1 Tf
(-strains)Tj
4.0875 0 Td
(\(they)Tj
2.262 0 Td
(exhibit)Tj
2.9764 0 Td
(up)Tj
1.2642 0 Td
(to)Tj
1.0262 0 Td
(65%)Tj
1.9275 0 Td
(of)Tj
1.0148 0 Td
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