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1.00964 (ions) -209.98200 (on) -209.01400 (the) ] TJ 11.95590 TL T* [ (pr) 44.98390 (oblem) -201.00900 (instances) -200.99200 (of) -201.01300 (J) 35.00890 (ug) -199.99600 (et) -200.98400 (al\056) -201.01100 (and) -201.01100 (scale) -201.01500 (to) -201.01300 (lar) 36.99140 (g) 10.00320 (er) -200.98400 (instances\054) ] TJ 11.95510 TL T* [ (le) 14.98100 (ver) 15.01470 (a) 10.00320 (ging) -320.00300 (mor) 14.99140 (al) -320.01300 (linea) 10.01790 (g) 10.00320 (e) -319.01000 (tr) 14.99140 (acing) -320.00700 (to) -320.01300 (pr) 14.99010 (actical) -320.00300 <7369676e690263616e6365> 15.00980 (\056) ] TJ /R44 11.95520 Tf 50.54690 TL T* [ (1\056) -249.99000 (Intr) 18.01460 (oduction) ] TJ /R46 9.96260 Tf 11.95510 -19.83790 Td [ (Recent) -302.01600 (adv) 24.98240 (ances) -301.01100 (in) -302.01600 (microscop) 9.98730 (y) -301.00400 (ha) 19.99790 (v) 14.00260 (e) -301.01300 (enabled) -302.01800 (biologists) ] TJ -11.95510 -11.95630 Td [ (to) -261.00300 (observ) 14.99140 (e) -261.02000 (or) 17.99880 (g) 4.98446 (anisms) -262.00300 (on) -261.00500 (a) -261.02000 (cellular) -261.00500 (le) 25.01790 (v) 14.98280 (el) -261.01300 (with) -262.01300 (higher) -261 (spatio\055) ] TJ 11.95510 TL T* [ (temporal) -357.01400 (resolution) -358.01700 (than) -356.98700 (before) -357.00400 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 185.53400 234.80000 Tm (12) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 195.49600 234.80000 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 201.55400 234.80000 Tm (16) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 211.51600 234.80000 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 217.56400 234.80000 Tm (44) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 227.52600 234.80000 Tm [ (\135\056) -632.01200 (Analysis) -357.98400 (of) ] TJ -177.41400 -11.95470 Td [ (such) -223.99000 (microscop) 9.98730 (y) -223.99700 (sequences) -223.99200 (is) -223.98800 (k) 10.00320 (e) 15.01220 (y) -223.99700 (to) -225.00900 (se) 25.01790 (v) 14.98280 (eral) -224.00700 (open) -223.99200 (questions) ] TJ 11.95510 TL T* [ (in) -370.00400 (biology) 65.00880 (\054) -400.98900 (including) -370.01400 (embryonic) -369.99000 (de) 25.01670 (v) 14.98280 (elopment) -369.98500 (of) -370.99400 (comple) 14.99750 (x) ] TJ T* [ (or) 17.99880 (g) 4.98446 (anisms) -224.00900 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 96.38870 198.93500 Tm (25) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 106.35100 198.93500 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 111.07300 198.93500 Tm (26) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 121.03600 198.93500 Tm [ (\135\054) -228.98300 (tissue) -222.99500 (formation) -224.01000 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 198.88400 198.93500 Tm (17) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 208.84600 198.93500 Tm [ (\135) -224.00400 (or) -224 (the) -223.99400 (understand\055) ] TJ -158.73400 -11.95510 Td [ (ing) -373.01900 (of) -372.99500 (metastati) 1.01454 (c) -373.00100 (beha) 19.99790 (vior) -372.98300 (of) -372.99500 (tumor) -373.01000 (cells) -372.00400 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 225.87200 186.98000 Tm (50) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 235.83500 186.98000 Tm [ (\135\056) -678.98500 (Ho) 24.98360 (we) 25.01300 (v) 14.98280 (er) 39.98600 (\054) ] TJ -185.72300 -11.95590 Td [ (to) -346.98900 (get) -346.01300 (from) -346.99100 (a) -345.98600 (sequence) -346.99700 (of) -346.01900 (ra) 15.00980 (w) -346.99800 (microscop) 9.98853 (y) -346.99600 (images) -345.99900 (to) -346.98900 (bio\055) ] TJ 11.95510 TL T* [ (logically) -224.00600 (or) -222.98100 (clinically) -224.00800 (rele) 25.01050 (v) 24.98110 (ant) -223.01400 (quantities\054) -229.01600 (such) -223.99000 (as) -222.98000 (cell) -223.99700 (motil\055) ] TJ T* [ (ity) 64.98920 (\054) -281.00900 (migration) -273.99100 (patterns) -274.99500 (and) -273.98700 (dif) 24.98480 (ferentiation) -274.98800 (schedules\054) -280.98700 (rob) 20.00400 (ust) ] TJ T* [ (methods) -192.99000 (for) ] TJ /R50 9.96260 Tf 49.21990 0 Td [ (cell) -192.98100 (linea) 10.01790 (g) 10.00320 (e) -192.01200 (tr) 14.99140 (acing) ] TJ /R46 9.96260 Tf 77.91020 0 Td [ (are) -193 (required) -192 (and) -192.98200 (ha) 19.99670 (v) 14.98280 (e) -192.99200 (there\055) ] TJ -127.13000 -11.95470 Td [ (fore) -250.00400 (recei) 25.00620 (v) 14.98280 (ed) -249.99700 (considerable) -250.01700 (attention) -250.00100 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 197.99700 127.20400 Tm (2) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 202.97900 127.20400 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 207.96000 127.20400 Tm (3) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 212.94100 127.20400 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 217.92200 127.20400 Tm (13) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 227.88500 127.20400 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 232.86600 127.20400 Tm (32) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 242.82900 127.20400 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 247.81000 127.20400 Tm (33) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 257.77200 127.20400 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 262.75400 127.20400 Tm (34) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 272.71600 127.20400 Tm (\135\056) Tj -210.64900 -12.86520 Td [ (Cell) -391.99000 (lineage) -392.00200 (tracing) -392.98100 (is) -392 (typically) -392.01700 (considered) -391.99600 (a) -392.99800 (t) 0.98758 (w) 10 (o) -392.98800 (step) ] TJ -11.95510 -11.95510 Td [ (problem\072) -400.01700 (In) -295.00800 (the) -293.98300 <02727374> -294.98500 (step\054) -306.01100 (indi) 25 (vidual) -294.98500 (cells) -294.99700 (are) -294.98300 (detected) -295.00200 (and) ] TJ ET Q 3.98000 w 0 G 501.12100 904.14800 m 1446.11000 904.14800 l S q 10 0 0 10 0 0 cm BT /R52 5.97760 Tf 1 0 0 1 60.14100 83.81290 Tm [ <03> -0.90058 ] TJ /R46 7.97010 Tf 4.31680 -2.81289 Td [ (Authors) -250.01000 (contrib) 19.99660 (uted) -250.01900 (equally) 64.97830 (\056) ] TJ ET Q q 2362.38000 0 0 1259.22000 3088.62000 4172.83000 cm /R56 Do Q q 10 0 0 10 0 0 cm BT /R46 8.96640 Tf 1 0 0 1 308.86200 406.32400 Tm [ (Figure) -377.02100 (1\056) -376.99300 (Depicted) -377.02100 (abo) 15.01100 (v) 14.98650 (e) -376.99100 (is) -377.00400 (a) -376.98800 (lineage) -377.01000 (forest) -377.00400 (of) -376.98800 (cells) -376.98800 (from) -375.99100 (a) -376.98800 (se\055) ] TJ 10.95820 TL T* [ (quence) -373.97900 (of) -373.02100 (microscop) 9.99010 (y) -374.00900 (images\056) -681.00900 (The) -372.98500 <02727374> -373.99300 (image) -372.98500 (of) -373.98200 (the) -373.01800 (sequence) ] TJ 10.95900 TL T* [ (is) -323.98200 (sho) 25.00930 (wn) -323.00500 (on) -323.99600 (the) -323.00500 (left\056) -531.01100 (The) -324.02100 (last) -323.99100 (image) -323.01600 (is) -323.98500 (sho) 25.01200 (wn) -323.00500 (on) -323.99600 (the) -324.00700 (right\056) -529.99000 (Cell) ] TJ T* [ (di) 25.01470 (visions) -249.99700 (are) -249.98700 (depicted) -250.01700 (in) -250.01400 (black\056) ] TJ /R46 9.96260 Tf 38.20310 TL T* [ (se) 15.01960 (gmented) -375.01100 (in) -374.00400 (e) 25.01050 (v) 14.00260 (er) 1.01209 (y) -374.99100 (image\056) -685.00300 (Then\054) -405.02100 (in) -374.98600 (the) -374.98600 (second) -374.98200 (step\054) -405.99600 (in\055) ] TJ 11.95470 TL T* [ (di) 24.98850 (vidual) -363.99300 (cells) -364.00500 (are) -365.01000 (track) 9.99833 (ed) -364.01800 (o) 14.98280 (v) 14.98280 (er) -363.98800 (time) -363.99300 (and\054) -392.98300 (in) -364.00800 (case) -363.98300 (of) -364.99800 (a) -363.98300 (cell) ] TJ T* [ (di) 24.98850 (vision\054) -257.99900 (link) 9.98363 (ed) -257.01600 (to) -255.98400 (their) -257.00400 (ancestor) -255.99200 (cell\054) -259.00600 (to) -255.98700 <026e616c6c79> -256.98200 (arri) 25.01050 (v) 14.98280 (e) -256.00100 (at) -257.01100 (the) ] TJ T* [ (lineage) -251.98500 (forest) -251.01000 (of) -251.99500 (all) -251.98700 (cells) -251.00500 (\050Fig\056) ] TJ ET Q 1 0 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 430.98400 299.38000 Tm (1) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 435.96500 299.38000 Tm [ (\051\056) -315.00600 (The) -251.99200 (tracking) -252.01700 (subproblem) ] TJ -127.10300 -11.95510 Td [ (is) -271.00100 (complicated) -270.98200 (by) -271.00400 (cells) -271.00100 (that) -270.99900 (enter) -271.01400 (or) -271.01400 (lea) 19.99670 (v) 14.98280 (e) -271.01800 (the) -271.98400 <02656c64> -270.98900 (of) -271.01400 (vie) 24.98600 (w) 65.01370 (\054) ] TJ 11.95590 TL T* [ (or) -303.00600 (lo) 24.98850 (w) -301.98600 (temporal) -302.98400 (resolution) -301.98900 (that) -302.99400 (allo) 24.99090 (ws) -302.01800 (lar) 17.99700 (ge) -303.00800 (displacements) ] TJ 11.95510 TL T* [ (or) -224 (e) 25.01050 (v) 14.98280 (en) -224.00200 (multiple) -223.99400 (consecuti) 25 (v) 14.98280 (e) -224.00700 (di) 24.98600 (visions) -224.00200 (within) -224.00700 (one) -223.99700 (time) -224.01700 (step\056) ] TJ T* [ (In) -307.98500 (addition) -309.01000 (to) -308.01500 (this\054) -323.01700 (mistak) 9.99343 (es) -307.98800 (made) -308.99700 (in) -308.01700 (the) -308.01700 <02727374> -308.98300 (step\054) -322.99000 (leading) ] TJ T* [ (to) -348.98800 (o) 14.98280 (v) 14.98280 (er) 19.98930 (\055) -349.00300 (or) -348.99800 (underse) 14.98770 (gmentation) -348.99600 (of) -348.02000 (the) -348.99100 (cells\054) -374.00100 (propag) 4.99426 (ate) -349 (into) ] TJ T* [ (the) -293.98300 (resulting) -293.99500 (lineage) -294.01700 (forest) -293.98000 (and) -293.98500 (cause) -295.01000 (spurious) -293.99500 (di) 24.98600 (visions) -293.99000 (or) ] TJ T* [ (missing) -255.00600 (branches\054) -255.98900 (respecti) 24.98600 (v) 14.98280 (ely) 65.01860 (\056) -323.98000 (The) -255.01100 (tracking) -254.99700 (subproblem) -255.01100 (is) ] TJ 11.95590 TL T* [ (closely) -304.00600 (related) -303.98600 (to) -304.01500 (multi\055t) 0.98268 (ar) 17.98720 (get) -303.97900 (tracking) -304.00800 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 469.90800 203.73800 Tm (11) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 479.87000 203.73800 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 485.38900 203.73800 Tm (42) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 495.35200 203.73800 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 500.87100 203.73800 Tm (47) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 510.83400 203.73800 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 516.35300 203.73800 Tm (20) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 526.31500 203.73800 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 531.83400 203.73800 Tm (43) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 541.79700 203.73800 Tm (\135) Tj -232.93500 -11.95510 Td [ (or) -245.01500 (reconstruc) 0.99003 (tion) -244.99300 (of) -245.01500 (tree\055lik) 10.00810 (e) -244.00300 (structures) -244.99800 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 469.78800 191.78300 Tm (15) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 479.75100 191.78300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 484.67200 191.78300 Tm (39) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 494.63500 191.78300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 499.56600 191.78300 Tm (37) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 509.52900 191.78300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 514.45000 191.78300 Tm (45) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 524.41300 191.78300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 529.34500 191.78300 Tm (46) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 539.30700 191.78300 Tm (\135\056) Tj -230.44500 -11.95510 Td [ (It) -291.00500 (has) -291.00300 (been) -291.00800 (cast) -291.00500 (in) -290.99800 (the) -291.00300 (form) -291 (of) -291.01000 (dif) 24.98600 (ferent) -291.01500 (optimization) -290.99100 (prob\055) ] TJ 11.95510 TL T* [ (lems) -387.99200 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 334.86400 167.87300 Tm (22) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 344.82700 167.87300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 351.19300 167.87300 Tm (24) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 361.15500 167.87300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 367.51200 167.87300 Tm (36) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 377.47400 167.87300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 383.84000 167.87300 Tm (40) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 393.80300 167.87300 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 400.15900 167.87300 Tm (41) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 410.12100 167.87300 Tm [ (\135) -388.01900 (that) -389.01800 (can) -388.01900 (deal) -388.98900 (with) -387.98900 (some) -387.99400 (of) -388.99400 (the) ] TJ -101.25900 -11.95510 Td [ (mentioned) -257.99900 (dif) 24.98600 <0263756c746965732c> -259.99300 (e\056g\056\054) -259.98600 (by) -257.98600 (selecting) -258.99100 (from) -257.98600 (multiple) -257.99100 (se) 15.01710 (g\055) ] TJ 11.95590 TL T* [ (mentation) -249.99700 (h) 4.98446 (ypotheses) -250.00500 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 401.23600 143.96200 Tm (40) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 411.19800 143.96200 Tm (\054) Tj ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 416.17900 143.96200 Tm (41) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 426.14200 143.96200 Tm (\135\056) Tj -105.32500 -15.14100 Td [ (Jug) -326.98700 (et) -327.00200 (al\056) -327 (\133) ] TJ ET Q 0 1 0 rg q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 364.62300 128.82100 Tm (21) Tj ET Q 0 g q 10 0 0 10 0 0 cm BT /R46 9.96260 Tf 1 0 0 1 374.58500 128.82100 Tm [ (\135\054) -347 (on) -326.99400 (the) -326.99700 (other) -326.99400 (hand\054) -346.00600 (ha) 19.99670 (v) 14.98280 (e) -327.00900 (proposed) -328 (a) -327.00900 (rig\055) ] TJ -65.72300 -11.95590 Td [ (orous) -292.99300 (mathematical) -293.01200 (abstraction) -292.00500 (of) -293.01000 (the) -293 (joint) -293.02000 (problem) -292.98500 (which) ] TJ 11.95510 TL T* [ (the) 14.98520 (y) -247.01300 (call) 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