author contributions: - eneuro sro\surs\ohqhfohdusodvwlffdjhv [ [ fp $oohqwrzq,qf 86$ zlwkfruqfre...

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Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreading process. This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution and reproduction in any medium provided that the original work is properly attributed. Copyright © 2016 Society for Neuroscience This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version. Research Article: New Research | Cognition and Behavior Effects of pre-pubertal or adult site-specific knockdown of estrogen receptor β in the medial preoptic area and medial amygdala on social behaviors in male mice Site-specific roles of ERβ in male social behavior Mariko Nakata 1 , Kazuhiro Sano 1 , Sergei Musatov 2,# , Naoko Yamaguchi 3 , Toshiro Sakamoto 4 and Sonoko Ogawa 1 1 Laboratory of Behavioral Neuroendocrinology, University of Tsukuba, Tsukuba, Ibaraki 305-8577, Japan 2 Laboratory of Molecular Neurosurgery, Weill Cornell University Medical College, New York 10021, USA 3 Department of Pharmacology, School of Medicine, Aichi Medical University, Nagakute, Aichi 480-1195, Japan 4 Department of Psychology, Kyoto Tachibana University, Kyoto 607-8175, Japan DOI: 10.1523/ENEURO.0155-15.2016 Received: 15 December 2015 Revised: 14 March 2016 Accepted: 15 March 2016 Published: 23 March 2016 Author Contributions: M.N., K.S., S.M., N.Y., T.S., and S.O. designed research; M.N., K.S. and N.Y. performed research; S.M. and S.O. contributed unpublished reagents/analytic tools; M.N. and K.S. analyzed data; M.N., K.S. and S.O. wrote the paper. Funding: Grant-in-aid for Scientific Research: #23240057. Grant-in-aid for Scientific Research: #15H05724. The Japan Society for Promotion of Science research: fellowship for young scientists DC2. Conflict of Interest: Authors report no conflict of interest. # S.M. was deceased on May 2015. Corresponding author: Sonoko Ogawa, PhD, Laboratory of Behavioral Neuroendocrinology, University of Tsukuba, Research Building D-409, 1-1-1 Tennodai, Tsukuba, Ibaraki 305-8577, Japan, E-mail: [email protected] Cite as: eNeuro 2016; 10.1523/ENEURO.0155-15.2016 Alerts: Sign up at eneuro.org/alerts to receive customized email alerts when the fully formatted version of this article is published.

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Page 1: Author Contributions: - eNeuro SRO\SURS\OHQHFOHDUSODVWLFFDJHV [ [ FP $OOHQWRZQ,QF 86$ ZLWKFRUQFRE 150 EHGGLQJ )RRGDQGZDWHUZHUHSURYLGHG ad libitum.$OOSURFHGXUHVZHUHFRQGXFWHGLQ 152 $QLPDO

Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreadingprocess.

This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International(http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution and reproduction in anymedium provided that the original work is properly attributed.

Copyright © 2016 Society for Neuroscience

This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version.

Research Article: New Research | Cognition and Behavior

Effects of pre-pubertal or adult site-specific knockdown of estrogen receptor β inthe medial preoptic area and medial amygdala on social behaviors in male mice

Site-specific roles of ERβ in male social behavior

Mariko Nakata1, Kazuhiro Sano1, Sergei Musatov2,#, Naoko Yamaguchi3, Toshiro Sakamoto4 and SonokoOgawa1

1Laboratory of Behavioral Neuroendocrinology, University of Tsukuba, Tsukuba, Ibaraki 305-8577, Japan2Laboratory of Molecular Neurosurgery, Weill Cornell University Medical College, New York 10021, USA3Department of Pharmacology, School of Medicine, Aichi Medical University, Nagakute, Aichi 480-1195, Japan4Department of Psychology, Kyoto Tachibana University, Kyoto 607-8175, Japan

DOI: 10.1523/ENEURO.0155-15.2016

Received: 15 December 2015

Revised: 14 March 2016

Accepted: 15 March 2016

Published: 23 March 2016

Author Contributions: M.N., K.S., S.M., N.Y., T.S., and S.O. designed research; M.N., K.S. and N.Y.performed research; S.M. and S.O. contributed unpublished reagents/analytic tools; M.N. and K.S. analyzeddata; M.N., K.S. and S.O. wrote the paper.

Funding: Grant-in-aid for Scientific Research: #23240057. Grant-in-aid for Scientific Research: #15H05724. TheJapan Society for Promotion of Science research: fellowship for young scientists DC2.

Conflict of Interest: Authors report no conflict of interest.

#S.M. was deceased on May 2015.

Corresponding author: Sonoko Ogawa, PhD, Laboratory of Behavioral Neuroendocrinology, Universityof Tsukuba, Research Building D-409, 1-1-1 Tennodai, Tsukuba, Ibaraki 305-8577, Japan, E-mail:[email protected]

Cite as: eNeuro 2016; 10.1523/ENEURO.0155-15.2016

Alerts: Sign up at eneuro.org/alerts to receive customized email alerts when the fully formatted version of thisarticle is published.

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16 17 18

19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39

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48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69

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70 71 72 73 74 75 76 77 78 79 80

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81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99

100 101 102 103 104 105 106 107 108 109 110 111 112

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113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142

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143 Experimental Animals 144

145 146 147 148 149

ad libitum.150 151 152 153 154 155

Estrogen Receptor Silencing using Small Hairpin RNA 156 157 158

Experiment 1 Experiments 2 159 and 3160

161 162 163 164 165 166 167 168 169 170 171 172 173

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174 175 176 177 178 179 180

Behavioral Tests 181 Sexual behavior test 182

183 184 185 186 187 188 189 190 191 192

Aggressive behavior test 193 194 195 196 197 198 199 200 201 202 203 204 205

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206 207

Sexual preference test 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224

sexual 225 behavior test 226

227 228 229 230 231 232 233 234 235 236

Quantitative analysis of behavioral data 237

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238 239 240 241 242 243 244 245

p 246 247

Histological Analysis248 Preparation of brain tissues for immunohistochemistry 249

250 251

i.p.252 253 254 255 256 257 258 259 260 261

Immunohistochemistry 262 263 264 265 266 267 268 269

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270 271 272 273 274 275 276 277 278 279 280 281 282 283

Analysis of immunopositive cells 284 285 286 287 288 289 290 291 292 293 294 295 296

p 297 298

Experimental Design 299 Experiment 1 Pre-pubertal treatment in the MPOA and MeA 300

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301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317

Experiment 2: Post-pubertal treatment in the MPOA 318 319 320 321 322

Experiment 1)323 324 325 326

Experiment 3: Post-pubertal treatment in the MeA 327 328 329 330 331 332

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333 334 335 336 337 338 339 340 341 342 343 344 345 346 347

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348 349

Effects of pre-pubertal silencing of ER in the MPOA on sexual and aggressive 350 behavior 351

352 353 354 355 356 357

F p358 F p359

n.s.360 361 362 363

β364 365 366 367 368 369

t p370 t p t p371

372 373

β374 375 376 377 378

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379 Effects of pre-pubertal silencing of ER in the MeA on sexual and aggressive 380 behaviors 381

382 383 384 385 386 387 388 389 390 391

t p t p392 t p393

β394 395 396 397 398 399 400 401 402 403 404 405

F p406 F p407

F p408 409 410

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411 412 413 414

F415 p F p416

417 418 419 420 421 422 423 424 425 426 427 428

t p429 t p430

n.s.431 432

t p t p433 434 435 436 437 438 439

F p F440 p F p441

442

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443 444 445 446 447

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448 449 450 451 452 453 454 455 456 457 458 459 460 461 462 463

Role of ER in the MPOA in the regulation of sexual and aggressive behavior 464 465 466 467 468 469 470 471 472 473 474 475 476 477 478

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479 480 481 482 483 484 485 486 487 488 489 490 491 492 493 494 495 496 497 498 499 500 501 502 503 504 505 506 507 508 509 510

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511 512 513 514 515 516 517 518 519

Role of ER in the MeA in the regulation of sexual and aggressive behavior 520 521 522 523 524 525 526 527 528 529 530 531 532 533 534 535 536 537 538 539 540 541 542

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543 544 545 546 547 548 549 550 551 552 553 554 555 556 557 558 559 560 561 562 563 564 565 566 567 568 569 570 571

572 573 Conclusion 574

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575 576 577 578 579 580 581 582 583

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584

585 586

587 588 589

590 591

592 593 594

595 596

597 598 599

600 601 602

603 604 605

606 607 608 609

610 611 612

613 614

615 616 617

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618 619 620

621 622 623 624

625 626

627 628

629 630 631

632 633 634

635 636 637 638

639 640 641

642 643

644 645 646

647 648 649

650 651 652

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653 654 655 656

657 658

659 660 661

662 663

664 665 666

667 668 669 670

671 672 673 674

675 676 677

678 679 680

681 682 683

684 685

686 687

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688 689 690

691 692

693 694 695

696 697 698 699

700 701 702

703 704 705

706 707 708 709

710 711

712 713 714

715 716

717 718 719

720 721

722

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723 724 725 726 727 728 729 730 731 732 733 734 735 736 737 738 739 740 741 742 743 744 745 746 747 748 749 750 751 752 753 754

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755 756 757 758 759 760 761 762 763 764 765 766 767 768 769 770 771 772 773 774 775 776 777 778 779 780 781 782 783 784 785 786

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787 788 789 790 791 792

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793 794 795 796 797 798 799 800 801 802 803 804

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p

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