%PDF-1.6
%
1 0 obj
<>
endobj
2 0 obj
<>stream
Katarína Bod’ová, Gabriel J. Mitchell, Roy Harpaz, Elad Schneidman, Gašper Tkačik
Probabilistic models of individual and collective animal behavior
www.plosone.org
www.plosone.org
endstream
endobj
3 0 obj
<>/XObject<>>>/CropBox[0 0 612 792]/MediaBox[0 0 612 792]/Parent 8 0 R/Annots 9 0 R/Contents 10 0 R/TrimBox[0 0 612 792]>>
endobj
9 0 obj
[11 0 R 12 0 R 13 0 R 14 0 R 15 0 R 16 0 R 17 0 R 18 0 R 19 0 R 20 0 R 21 0 R 22 0 R 23 0 R 24 0 R 25 0 R 26 0 R 27 0 R 28 0 R 29 0 R 30 0 R 31 0 R]
endobj
11 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref001)>>
endobj
12 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref008)>>
endobj
13 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref001)>>
endobj
14 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref004)>>
endobj
15 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref005)>>
endobj
16 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref009)>>
endobj
17 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref011)>>
endobj
18 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref012)>>
endobj
19 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref013)>>
endobj
20 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref015)>>
endobj
21 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref016)>>
endobj
22 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref017)>>
endobj
23 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref018)>>
endobj
24 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref019)>>
endobj
25 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref020)>>
endobj
26 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref021)>>
endobj
27 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref022)>>
endobj
28 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref023)>>
endobj
29 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref024)>>
endobj
30 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref025)>>
endobj
31 0 obj
<>/Border[0 0 0]/A 32 0 R>>
endobj
32 0 obj
<>
endobj
10 0 obj
[33 0 R 34 0 R 35 0 R 36 0 R 37 0 R 38 0 R 39 0 R 40 0 R 41 0 R 42 0 R 43 0 R 44 0 R 45 0 R 46 0 R 47 0 R 48 0 R 49 0 R 50 0 R 51 0 R 52 0 R]
endobj
33 0 obj
<>stream
q
0.83 0.64 0.02 0 k
265.7764 628.3276 m
270.5386 628.3276 l
h
f*
275.6976 628.3276 m
280.4598 628.3276 l
h
f*
251.0929 589.3228 m
255.8551 589.3228 l
h
f*
260.2772 589.3228 m
265.0394 589.3228 l
h
f*
269.5181 589.3228 m
274.2803 589.3228 l
h
f*
463.2378 524.2961 m
468 524.2961 l
h
f*
473.1591 524.2961 m
482.6835 524.2961 l
h
f*
215.2063 472.3087 m
224.7307 472.3087 l
h
f*
461.9339 459.326 m
471.4583 459.326 l
h
f*
476.6173 459.326 m
486.1417 459.326 l
h
f*
461.7071 446.3433 m
471.1748 446.3433 l
h
f*
312.7748 433.3039 m
322.2992 433.3039 l
h
f*
326.778 433.3039 m
336.3024 433.3039 l
h
f*
467.0929 433.3039 m
476.6173 433.3039 l
h
f*
395.1496 420.3213 m
404.674 420.3213 l
h
f*
357.7323 212.315 m
367.2567 212.315 l
h
f*
393.5622 212.315 m
403.0866 212.315 l
h
f*
358.8094 186.3496 m
368.3339 186.3496 l
h
f*
379.6157 186.3496 m
389.0835 186.3496 l
h
f*
417.6567 186.3496 m
427.1811 186.3496 l
h
f*
0 g
1 j
1 J
0 w
10 0 0 10 200.0125 707.4141 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(individual)Tj
4.2859 0 Td
(behavior)Tj
3.7134 0 Td
(on)Tj
1.2756 0 Td
(a)Tj
0.6519 0 Td
(moment-by-moment)Tj
8.7193 0 Td
(basis)Tj
2.1487 0 Td
(and)Tj
1.7291 0 Td
(what)Tj
2.1657 0 Td
(factors)Tj
2.9026 0 Td
(influence)Tj
3.9175 0 Td
(behavioral)Tj
-31.5097 -1.3039 Td
(decisions?)Tj
4.2746 0 Td
(How)Tj
2.1826 0 Td
(can)Tj
1.6214 0 Td
(coordinated)Tj
5.057 0 Td
(and)Tj
1.7291 0 Td
(collective)Tj
3.9345 0 Td
(motion)Tj
3.1691 0 Td
(realistically)Tj
4.6942 0 Td
(emerge)Tj
3.1464 0 Td
(in)Tj
1.0318 0 Td
(groups)Tj
2.9707 0 Td
(of)Tj
1.0148 0 Td
(inter-)Tj
-34.8262 -1.2982 Td
(acting)Tj
2.6532 0 Td
(animals?)Tj
3.702 0 Td
(How)Tj
2.1826 0 Td
(do)Tj
1.253 0 Td
(we)Tj
1.3266 0 Td
(include)Tj
3.1861 0 Td
(the)Tj
1.4683 0 Td
(existence)Tj
3.8551 0 Td
(of)Tj
1.0148 0 Td
(different)Tj
3.6567 0 Td
(cognitive)Tj
4.6601 0 Td
(or)Tj
1.0942 0 Td
(behavioral)Tj
4.3936 0 Td
(states)Tj
-34.4463 -1.2983 Td
(into)Tj
1.8425 0 Td
(mathematical)Tj
5.6409 0 Td
(descriptions)Tj
5.0966 0 Td
(of)Tj
1.0148 0 Td
(animal)Tj
2.9707 0 Td
(motion?)Tj
-15.3693 -1.2983 Td
(Recent)Tj
2.9593 0 Td
(progress)Tj
3.6 0 Td
(in)Tj
1.0375 0 Td
(automated)Tj
4.49 0 Td
(recording)Tj
4.1216 0 Td
(techniques)Tj
4.5297 0 Td
(opened)Tj
3.1691 0 Td
(the)Tj
1.4684 0 Td
(door)Tj
2.1316 0 Td
(to)Tj
1.0261 0 Td
(addressing)Tj
4.5071 0 Td
(these)Tj
-34.2366 -1.3039 Td
(questions,)Tj
4.2462 0 Td
(by)Tj
1.1792 0 Td
(making)Tj
3.2542 0 Td
(it)Tj
0.7823 0 Td
(possible)Tj
3.4129 0 Td
(to)Tj
1.0205 0 Td
(track)Tj
2.245 0 Td
(single)Tj
2.5342 0 Td
(or)Tj
1.0998 0 Td
(multiple)Tj
3.566 0 Td
(interacting)Tj
4.5637 0 Td
(animals)Tj
3.3279 0 Td
(for)Tj
1.3889 0 Td
(extended)Tj
-32.6208 -1.2982 Td
(periods)Tj
3.1917 0 Td
(of)Tj
1.0148 0 Td
(time)Tj
2.0296 0 Td
([)Tj
0.83 0.64 0.02 0 k
(1)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(8)Tj
0 g
(].)Tj
2.5966 0 Td
(Traditional)Tj
4.7281 0 Td
(models)Tj
3.1068 0 Td
(of)Tj
1.0148 0 Td
(animal)Tj
2.9706 0 Td
(motion,)Tj
3.3903 0 Td
(constructed)Tj
4.9322 0 Td
(to)Tj
1.0262 0 Td
(favor)Tj
2.2733 0 Td
(simplicity)Tj
-32.275 -1.2983 Td
(and)Tj
1.7291 0 Td
(provide)Tj
3.2881 0 Td
(a)Tj
0.652 0 Td
(qualitative)Tj
4.3823 0 Td
(match)Tj
2.7213 0 Td
(to)Tj
1.0261 0 Td
(the)Tj
1.4684 0 Td
(observed)Tj
3.7984 0 Td
(collective)Tj
3.9288 0 Td
(or)Tj
1.0998 0 Td
(aggregate)Tj
4.0025 0 Td
(\(averaged\))Tj
4.4333 0 Td
(behaviors,)Tj
-32.5301 -1.3039 Td
(are)Tj
1.4456 0 Td
(today)Tj
2.4435 0 Td
(being)Tj
2.4321 0 Td
(revisited)Tj
3.6113 0 Td
(and)Tj
1.7348 0 Td
(carefully)Tj
3.634 0 Td
(fitted)Tj
2.3244 0 Td
(to)Tj
1.0204 0 Td
(extensive)Tj
3.8891 0 Td
(data)Tj
1.9106 0 Td
(to)Tj
1.0261 0 Td
(provide)Tj
3.2882 0 Td
(predictive,)Tj
4.422 0 Td
(quantita-)Tj
-33.1821 -1.2983 Td
(tive)Tj
1.661 0 Td
(models)Tj
3.1068 0 Td
([)Tj
0.83 0.64 0.02 0 k
(1)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
1.2586 0 Td
(4)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
0.9241 0 Td
(5)Tj
0 g
(].)Tj
-5.7543 -1.2983 Td
(In)Tj
1.1111 0 Td
(addition)Tj
3.6 0 Td
(to)Tj
1.0205 0 Td
(revisiting)Tj
3.9458 0 Td
(existing)Tj
3.3165 0 Td
(models,)Tj
3.3278 0 Td
(the)Tj
1.4741 0 Td
(wealth)Tj
2.8346 0 Td
(of)Tj
1.0148 0 Td
(data)Tj
1.9105 0 Td
(also)Tj
1.7688 0 Td
(provides)Tj
3.651 0 Td
(the)Tj
1.4684 0 Td
(motivation)Tj
4.6204 0 Td
(to)Tj
-36.2605 -1.3039 Td
(devise)Tj
2.6758 0 Td
(new)Tj
1.8766 0 Td
(models)Tj
3.1067 0 Td
(and)Tj
1.7291 0 Td
(remove)Tj
3.2202 0 Td
(some)Tj
2.33 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(previously-made)Tj
6.9051 0 Td
(restrictions)Tj
4.7225 0 Td
(and)Tj
1.7291 0 Td
(assumptions,)Tj
5.4482 0 Td
(or)Tj
-36.2265 -1.2982 Td
(at)Tj
0.9524 0 Td
(least)Tj
1.9842 0 Td
(test)Tj
1.5988 0 Td
(rigorously)Tj
4.2973 0 Td
(whether)Tj
3.4752 0 Td
(they)Tj
1.9276 0 Td
(are)Tj
1.4456 0 Td
(essential.)Tj
3.8098 0 Td
(A)Tj
0.907 0 Td
(central)Tj
2.9651 0 Td
(assumption)Tj
4.8585 0 Td
(that)Tj
1.7802 0 Td
(underlies)Tj
3.9174 0 Td
(many)Tj
-33.9191 -1.2983 Td
(models)Tj
3.101 0 Td
(of)Tj
1.0148 0 Td
(animal)Tj
2.9707 0 Td
(motion)Tj
3.1691 0 Td
(and)Tj
1.7292 0 Td
(behavior)Tj
3.7133 0 Td
(is)Tj
0.8504 0 Td
(that)Tj
1.7745 0 Td
(each)Tj
2.0296 0 Td
(animal)Tj
2.9707 0 Td
(continuously)Tj
5.4368 0 Td
(performs)Tj
3.8778 0 Td
(a)Tj
0.6576 0 Td
(universal)Tj
-33.2955 -1.3039 Td
(computation)Tj
5.3687 0 Td
(to)Tj
1.0262 0 Td
(determine)Tj
4.3199 0 Td
(its)Tj
1.1452 0 Td
(next)Tj
1.9559 0 Td
(action)Tj
2.6986 0 Td
(or)Tj
1.0998 0 Td
(movement)Tj
4.5128 0 Td
(direction)Tj
3.855 0 Td
([)Tj
0.83 0.64 0.02 0 k
(9)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(11)Tj
0 g
(].)Tj
3.0728 0 Td
(Animals,)Tj
3.8041 0 Td
(however,)Tj
-32.859 -1.2983 Td
(often)Tj
2.2847 0 Td
(exhibit)Tj
2.9763 0 Td
(discrete)Tj
3.3165 0 Td
(stereotyped)Tj
4.8246 0 Td
(behaviors)Tj
4.0705 0 Td
(\(termed)Tj
3.4242 0 Td
(here)Tj
1.9673 0 Td
(behavioral)Tj
4.3936 0 Td
(states\))Tj
2.7383 0 Td
(or)Tj
1.0941 0 Td
(stereotyped)Tj
-31.0901 -1.2982 Td
(switches)Tj
3.5659 0 Td
(between)Tj
3.5263 0 Td
(these)Tj
2.2564 0 Td
(states)Tj
2.3924 0 Td
(that)Tj
1.7801 0 Td
(result)Tj
2.4491 0 Td
(in)Tj
1.0375 0 Td
(apparently)Tj
4.4617 0 Td
(very)Tj
1.9276 0 Td
(different)Tj
3.6566 0 Td
(modes)Tj
2.8573 0 Td
(of)Tj
1.0148 0 Td
(motion)Tj
3.1692 0 Td
(through)Tj
-34.0949 -1.2983 Td
(space,)Tj
2.6021 0 Td
(as)Tj
1.0148 0 Td
(for)Tj
1.389 0 Td
(example)Tj
3.549 0 Td
(kinematic)Tj
4.5864 0 Td
(proxies)Tj
3.5263 0 Td
(for)Tj
1.3889 0 Td
(different)Tj
3.6567 0 Td
(cognitive)Tj
3.8721 0 Td
(or)Tj
1.0998 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3924 0 Td
(described)Tj
-33.4712 -1.3039 Td
(by)Tj
1.1792 0 Td
([)Tj
0.83 0.64 0.02 0 k
(12)Tj
0 g
(].)Tj
2.0806 0 Td
(These)Tj
2.5625 0 Td
(individual)Tj
4.2916 0 Td
(animal)Tj
2.965 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3981 0 Td
(can)Tj
1.6271 0 Td
(be)Tj
1.1508 0 Td
(extracted)Tj
3.8835 0 Td
(from)Tj
2.2053 0 Td
(data)Tj
1.9162 0 Td
(using)Tj
2.3868 0 Td
(various)Tj
-33.0404 -1.2983 Td
(approaches:)Tj
4.9605 0 Td
(low-dimensional)Tj
6.9562 0 Td
(reduction)Tj
4.1159 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(complex)Tj
3.6227 0 Td
(behavior)Tj
3.7134 0 Td
([)Tj
0.83 0.64 0.02 0 k
(13)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(15)Tj
0 g
(],)Tj
3.5432 0 Td
(fragmentation)Tj
5.9244 0 Td
(of)Tj
-35.3194 -1.2983 Td
(trajectories)Tj
4.6544 0 Td
(into)Tj
1.8425 0 Td
(segments)Tj
3.9175 0 Td
(based)Tj
2.4718 0 Td
(on)Tj
1.2756 0 Td
(the)Tj
1.474 0 Td
(functionality)Tj
5.3291 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(space)Tj
2.3811 0 Td
([)Tj
0.83 0.64 0.02 0 k
(16)Tj
0 g
(],)Tj
2.0749 0 Td
(machine)Tj
3.6624 0 Td
(learning-based)Tj
-31.5664 -1.3039 Td
(detection)Tj
3.9344 0 Td
(of)Tj
1.0148 0 Td
(behavior)Tj
3.7134 0 Td
(types)Tj
2.2733 0 Td
([)Tj
0.83 0.64 0.02 0 k
(17)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
1.7405 0 Td
(18)Tj
0 g
(])Tj
1.508 0 Td
(or)Tj
1.0999 0 Td
(other)Tj
2.347 0 Td
(data)Tj
1.9162 0 Td
(mining)Tj
3.1351 0 Td
(methods)Tj
3.6851 0 Td
([)Tj
0.83 0.64 0.02 0 k
(19)Tj
0 g
(].)Tj
2.0749 0 Td
(Moreover,)Tj
4.3993 0 Td
(the)Tj
1.4684 0 Td
(states)Tj
-34.3103 -1.2982 Td
(can)Tj
1.627 0 Td
(be)Tj
1.1509 0 Td
(defined)Tj
3.2258 0 Td
(also)Tj
1.7631 0 Td
(for)Tj
1.389 0 Td
(the)Tj
1.4683 0 Td
(whole)Tj
2.6079 0 Td
(animal)Tj
2.9707 0 Td
(groups)Tj
2.9707 0 Td
([)Tj
0.83 0.64 0.02 0 k
(20)Tj
0 g
(],)Tj
2.0749 0 Td
(which)Tj
2.6532 0 Td
(we)Tj
1.3266 0 Td
(do)Tj
1.2529 0 Td
(not)Tj
1.5761 0 Td
(consider)Tj
3.6453 0 Td
(here.)Tj
2.194 0 Td
(The)Tj
-33.8964 -1.2983 Td
(switches)Tj
3.5659 0 Td
(between)Tj
3.5263 0 Td
(the)Tj
1.4683 0 Td
(individual's)Tj
4.8642 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3981 0 Td
(seem)Tj
2.245 0 Td
(to)Tj
1.0262 0 Td
(occur)Tj
2.4718 0 Td
(stochastically,)Tj
5.7316 0 Td
(but)Tj
1.5534 0 Td
(their)Tj
2.1033 0 Td
(rate)Tj
-35.3478 -1.3039 Td
(could)Tj
2.4491 0 Td
(in)Tj
1.0374 0 Td
(principle)Tj
3.8041 0 Td
(be)Tj
1.1509 0 Td
(affected)Tj
3.3222 0 Td
(by)Tj
1.1848 0 Td
(many)Tj
2.4775 0 Td
(factors:)Tj
3.1238 0 Td
(by)Tj
1.1848 0 Td
(spatial)Tj
2.778 0 Td
(preferences,)Tj
5.0173 0 Td
(by)Tj
1.1848 0 Td
(various)Tj
3.1408 0 Td
(environmen-)Tj
-31.8555 -1.2983 Td
(tal)Tj
1.1962 0 Td
(signals,)Tj
3.1294 0 Td
(by)Tj
1.1792 0 Td
(the)Tj
1.474 0 Td
(motion)Tj
3.1634 0 Td
(of)Tj
1.0148 0 Td
(conspecifics)Tj
5.0343 0 Td
(within)Tj
2.812 0 Td
(the)Tj
1.4683 0 Td
(group,)Tj
2.8347 0 Td
(etc.)Tj
1.593 0 Td
(The)Tj
1.7802 0 Td
(question)Tj
3.6623 0 Td
(therefore)Tj
3.8494 0 Td
(arises)Tj
-34.1912 -1.2983 Td
(about)Tj
2.4944 0 Td
(how)Tj
1.9389 0 Td
(to)Tj
1.0205 0 Td
(combine)Tj
3.719 0 Td
(in)Tj
1.0375 0 Td
(a)Tj
0.6519 0 Td
(single)Tj
2.5342 0 Td
(tractable)Tj
3.651 0 Td
(model,)Tj
2.9707 0 Td
(on)Tj
1.2756 0 Td
(the)Tj
1.4683 0 Td
(one)Tj
1.7008 0 Td
(hand,)Tj
2.4831 0 Td
(the)Tj
1.4684 0 Td
(kinematic)Tj
4.2009 0 Td
(description)Tj
-32.6152 -1.2982 Td
(of)Tj
1.0147 0 Td
(individual's)Tj
4.8643 0 Td
(motion)Tj
3.1634 0 Td
(through)Tj
3.4469 0 Td
(space,)Tj
2.6079 0 Td
(and)Tj
1.7291 0 Td
(on)Tj
1.2756 0 Td
(the)Tj
1.4683 0 Td
(other,)Tj
2.5739 0 Td
(the)Tj
1.4683 0 Td
(existence)Tj
3.8551 0 Td
(of)Tj
1.0148 0 Td
(different)Tj
3.6567 0 Td
(discrete)Tj
-32.139 -1.3039 Td
(behaviors,)Tj
4.2972 0 Td
(i.e.,)Tj
1.5874 0 Td
(behavioral)Tj
4.3937 0 Td
(states.)Tj
2.6192 0 Td
(If)Tj
0.8504 0 Td
(such)Tj
2.0579 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3924 0 Td
(are)Tj
1.4514 0 Td
(indeed)Tj
2.9366 0 Td
(present)Tj
3.1635 0 Td
(and)Tj
1.7291 0 Td
(important,)Tj
-31.8725 -1.2983 Td
(then)Tj
2.0182 0 Td
(ignoring)Tj
3.6567 0 Td
(their)Tj
2.1089 0 Td
(existencewhich)Tj
7.2113 0 Td
(happens)Tj
3.549 0 Td
(in,)Tj
1.2586 0 Td
(e.g.,)Tj
1.7858 0 Td
(classic)Tj
2.7382 0 Td
(zonal)Tj
2.3641 0 Td
(or)Tj
1.0998 0 Td
(force-field)Tj
4.337 0 Td
(modelswill)Tj
-32.1276 -1.2983 Td
(enable)Tj
2.8006 0 Td
(us)Tj
1.1111 0 Td
(to)Tj
1.0262 0 Td
(learn)Tj
2.2393 0 Td
(only)Tj
1.9786 0 Td
(laws)Tj
1.9446 0 Td
(of)Tj
1.0147 0 Td
(behavior)Tj
3.7134 0 Td
(and)Tj
1.7291 0 Td
(motion)Tj
3.1691 0 Td
(that)Tj
1.7745 0 Td
(are)Tj
1.4514 0 Td
(averaged)Tj
4.1442 0 Td
(over)Tj
2.3697 0 Td
(various)Tj
3.1408 0 Td
(behav-)Tj
-33.6073 -1.3039 Td
(ioral)Tj
2.0466 0 Td
(states.)Tj
2.6191 0 Td
(We)Tj
1.6101 0 Td
(would)Tj
2.7099 0 Td
(thus)Tj
1.9389 0 Td
(fail)Tj
1.4513 0 Td
(to)Tj
1.0262 0 Td
(capture)Tj
3.2144 0 Td
(the)Tj
1.474 0 Td
(observed)Tj
3.7928 0 Td
(heterogeneity)Tj
5.6522 0 Td
(due)Tj
1.7008 0 Td
(to)Tj
1.0261 0 Td
(the)Tj
1.4684 0 Td
(stochastic)Tj
4.1215 0 Td
(and)Tj
-35.8523 -1.2982 Td
(discrete)Tj
3.3165 0 Td
(nature)Tj
2.8233 0 Td
(of)Tj
1.0148 0 Td
(individual)Tj
4.2859 0 Td
(animal)Tj
2.9707 0 Td
(behaviors)Tj
4.0705 0 Td
(and,)Tj
1.9559 0 Td
(consequently,)Tj
5.7146 0 Td
(motion)Tj
3.1691 0 Td
(trajectories.)Tj
-28.1251 -1.2983 Td
(The)Tj
1.7858 0 Td
(combination)Tj
5.3461 0 Td
(of)Tj
1.0148 0 Td
(stochastic)Tj
4.1215 0 Td
(state)Tj
2.0353 0 Td
(transitions)Tj
4.4844 0 Td
(with)Tj
1.9956 0 Td
(deterministic)Tj
5.5161 0 Td
(laws)Tj
1.9446 0 Td
(of)Tj
1.0148 0 Td
(motion)Tj
3.1691 0 Td
(through)Tj
-33.6243 -1.3039 Td
(space)Tj
2.3754 0 Td
(distinguishes)Tj
5.4254 0 Td
(our)Tj
1.6271 0 Td
(model)Tj
2.7439 0 Td
(from)Tj
2.2054 0 Td
(the)Tj
1.4683 0 Td
(classical)Tj
3.4186 0 Td
(models)Tj
3.1011 0 Td
(where)Tj
2.6532 0 Td
(behavioral)Tj
4.3936 0 Td
(rules)Tj
2.1487 0 Td
(are)Tj
1.4457 0 Td
(assumed)Tj
-33.0064 -1.2983 Td
(fixed)Tj
2.1883 0 Td
(in)Tj
1.0375 0 Td
(time.)Tj
2.2563 0 Td
(Traditionally,)Tj
5.6579 0 Td
(such)Tj
2.058 0 Td
(rules)Tj
2.1486 0 Td
(are)Tj
1.4514 0 Td
(inferred)Tj
3.4412 0 Td
(by)Tj
1.1792 0 Td
(fitting)Tj
2.6532 0 Td
(to)Tj
1.0261 0 Td
(all)Tj
1.1452 0 Td
(available)Tj
3.668 0 Td
(data,)Tj
2.1374 0 Td
(resulting)Tj
3.7303 0 Td
(in)Tj
-35.7786 -1.2983 Td
(simulated)Tj
4.1215 0 Td
(trajectories)Tj
4.6601 0 Td
(which)Tj
2.6476 0 Td
(tend)Tj
2.0182 0 Td
(to)Tj
1.0262 0 Td
(be)Tj
1.1508 0 Td
(much)Tj
2.5172 0 Td
(smoother)Tj
4.0365 0 Td
(than)Tj
2.0296 0 Td
(real)Tj
1.6951 0 Td
(data.)Tj
2.1373 0 Td
(Our)Tj
1.8652 0 Td
(approach)Tj
3.9514 0 Td
(also)Tj
1.7689 0 Td
(dif-)Tj
-35.6256 -1.3039 Td
(fers)Tj
1.661 0 Td
(from)Tj
2.2054 0 Td
(the)Tj
1.474 0 Td
(population)Tj
4.5864 0 Td
(dynamics)Tj
4.0592 0 Td
(models)Tj
3.1011 0 Td
(capturing)Tj
4.0762 0 Td
(proportions)Tj
4.9946 0 Td
(of)Tj
1.0148 0 Td
(animals)Tj
3.3278 0 Td
(with)Tj
1.9956 0 Td
(different)Tj
-32.4961 -1.2982 Td
(tasks)Tj
2.1713 0 Td
(or)Tj
1.0941 0 Td
(behavioral)Tj
4.3937 0 Td
(states.)Tj
2.6249 0 Td
(For)Tj
1.6214 0 Td
(instance)Tj
3.5263 0 Td
([)Tj
0.83 0.64 0.02 0 k
(21)Tj
0 g
(])Tj
1.8538 0 Td
(and)Tj
1.7291 0 Td
([)Tj
0.83 0.64 0.02 0 k
(22)Tj
0 g
(])Tj
1.8482 0 Td
(study)Tj
2.3924 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3924 0 Td
(in)Tj
1.0375 0 Td
(honeybees)Tj
-31.0788 -1.2983 Td
(when)Tj
2.4037 0 Td
(multiple)Tj
3.566 0 Td
(food)Tj
2.0466 0 Td
(sources)Tj
3.1974 0 Td
(are)Tj
1.4457 0 Td
(present)Tj
3.1634 0 Td
(and)Tj
1.7348 0 Td
(distinguish)Tj
4.6375 0 Td
(between)Tj
3.5263 0 Td
(proportion)Tj
4.6317 0 Td
(of)Tj
1.0148 0 Td
(population)Tj
-31.3679 -1.2983 Td
(feeding)Tj
3.1634 0 Td
(on)Tj
1.2756 0 Td
(different)Tj
3.6566 0 Td
(sources.)Tj
3.4186 0 Td
(Similarly,)Tj
4.0252 0 Td
([)Tj
0.83 0.64 0.02 0 k
(23)Tj
0 g
(],)Tj
2.0806 0 Td
([)Tj
0.83 0.64 0.02 0 k
(24)Tj
0 g
(],)Tj
2.0749 0 Td
(and)Tj
1.7292 0 Td
([)Tj
0.83 0.64 0.02 0 k
(25)Tj
0 g
(])Tj
1.8538 0 Td
(distinguish)Tj
4.6375 0 Td
(behavioral)Tj
4.3936 0 Td
(states)Tj
2.3925 0 Td
(in)Tj
1.0374 0 Td
(ant)Tj
-35.7389 -1.3039 Td
(foraging)Tj
3.5659 0 Td
(due)Tj
1.7008 0 Td
(to)Tj
1.0205 0 Td
(either)Tj
2.5341 0 Td
(multiple)Tj
3.566 0 Td
(food)Tj
2.0466 0 Td
(sources)Tj
3.1974 0 Td
(or)Tj
1.0999 0 Td
(due)Tj
1.6951 0 Td
(to)Tj
1.0261 0 Td
(recruiting)Tj
4.1839 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(resting)Tj
2.9593 0 Td
(ants.)Tj
2.0863 0 Td
(While)Tj
-33.1651 -1.2983 Td
(these)Tj
2.2506 0 Td
(works)Tj
2.6419 0 Td
(describe)Tj
3.5263 0 Td
(different)Tj
3.6567 0 Td
(behavioral)Tj
4.7848 0 Td
(states,)Tj
3.0104 0 Td
(they)Tj
1.9275 0 Td
(do)Tj
1.2529 0 Td
(so)Tj
1.0885 0 Td
(in)Tj
1.0318 0 Td
(terms)Tj
2.4945 0 Td
(of)Tj
1.0148 0 Td
(population)Tj
4.5921 0 Td
(averaged)Tj
-33.2728 -1.2982 Td
(quantities,)Tj
4.3766 0 Td
(while)Tj
2.3698 0 Td
(our)Tj
1.627 0 Td
(goal)Tj
1.8709 0 Td
(is)Tj
0.8447 0 Td
(to)Tj
1.0261 0 Td
(incorporate)Tj
4.8926 0 Td
(a)Tj
0.652 0 Td
(set)Tj
1.3039 0 Td
(of)Tj
1.0148 0 Td
(different)Tj
3.6567 0 Td
(behaviors)Tj
4.0705 0 Td
(at)Tj
0.9524 0 Td
(the)Tj
1.4684 0 Td
(individual)Tj
4.2916 0 Td
(level)Tj
-34.418 -1.3039 Td
(and)Tj
1.7291 0 Td
(fit)Tj
1.0715 0 Td
(that)Tj
1.7801 0 Td
(to)Tj
1.0261 0 Td
(individual)Tj
4.286 0 Td
(trajectory)Tj
4.0592 0 Td
(data.)Tj
-12.7558 -1.2983 Td
(We)Tj
1.61 0 Td
(propose)Tj
3.4243 0 Td
(a)Tj
0.6576 0 Td
(class)Tj
2.0409 0 Td
(of)Tj
1.0148 0 Td
(probabilistic)Tj
5.1987 0 Td
(models)Tj
3.1011 0 Td
(that)Tj
1.7802 0 Td
(is)Tj
0.8447 0 Td
(sufficiently)Tj
4.6148 0 Td
(rich)Tj
1.8141 0 Td
(to)Tj
1.0205 0 Td
(capture)Tj
3.2201 0 Td
(a)Tj
0.652 0 Td
(wide)Tj
2.1203 0 Td
(variety)Tj
-34.3103 -1.2983 Td
(of)Tj
1.0091 0 Td
(complex)Tj
3.617 0 Td
(individual)Tj
4.2803 0 Td
(and)Tj
1.7234 0 Td
(collective)Tj
3.9288 0 Td
(behaviors,)Tj
4.2916 0 Td
(and)Tj
1.7291 0 Td
(very)Tj
1.9163 0 Td
(flexible)Tj
3.0784 0 Td
(in)Tj
1.0318 0 Td
(how)Tj
1.9332 0 Td
(external)Tj
3.4242 0 Td
(factors)Tj
2.897 0 Td
(modu-)Tj
-34.8602 -1.3039 Td
(late)Tj
1.6214 0 Td
(the)Tj
1.4683 0 Td
(focal)Tj
2.1203 0 Td
(animal's)Tj
3.5433 0 Td
(behavior.)Tj
3.9401 0 Td
(Despite)Tj
3.2428 0 Td
(this)Tj
1.6781 0 Td
(expressive)Tj
4.286 0 Td
(power,)Tj
2.948 0 Td
(the)Tj
1.4683 0 Td
(proposed)Tj
3.9572 0 Td
(models)Tj
3.1011 0 Td
(remain)Tj
-33.3749 -1.2982 Td
(easy)Tj
1.8935 0 Td
(to)Tj
1.0261 0 Td
(simulate)Tj
3.5943 0 Td
(given)Tj
2.3811 0 Td
(the)Tj
1.4683 0 Td
(parameters,)Tj
4.8983 0 Td
(and)Tj
1.7291 0 Td
(support)Tj
3.3222 0 Td
(tractable)Tj
3.6567 0 Td
(maximum)Tj
4.371 0 Td
(likelihood)Tj
4.2349 0 Td
(parameter)Tj
ET
Q
q
1 j
1 J
0 w
576 737.1 m
36 737.1 l
36 737.6 l
576 737.6 l
f*
36 741.2598 107.1496 23.6976 re
W* n
q
107.0929 0 0 23.6409 36 741.3165 cm
q
/I0 Do
Q
Q
Q
q
0 0 612 792 re
W* n
1 j
1 J
0 w
7.9999 0 0 7.9999 441.5811 745.7952 cm
BT
/F0 1 Tf
1 TL
-0.005 Tc
0 0 Td
(Probabilist)Tj
4.5709 0 Td
(ic)Tj
0.9283 0 Td
(models)Tj
3.3874 0 Td
(of)Tj
1.0418 0 Td
(animal)Tj
3.111 0 Td
(behavio)Tj
3.43 0 Td
(r)Tj
ET
Q
q
1 j
1 J
0 w
36 48.0002 m
576 48.0002 l
576 47.5002 l
36 47.5002 l
f*
0.83 0.64 0.02 0 k
81.5811 34.9228 m
237.8835 34.9228 l
h
f*
0 g
7.9999 0 0 7.9999 36 36 cm
BT
/F0 1 Tf
1 TL
-0.005 Tc
0 0 Td
(PLOS)Tj
2.8559 0 Td
(ONE)Tj
2.3669 0 Td
(|)Tj
0.83 0.64 0.02 0 k
0.4748 0 Td
(https://doi.or)Tj
5.4142 0 Td
(g/10.137)Tj
3.8198 0 Td
(1/journal.po)Tj
5.1236 0 Td
(ne.01930)Tj
4.089 0 Td
(49)Tj
0 g
2.0906 0 Td
(March)Tj
2.9622 0 Td
(7,)Tj
1.0418 0 Td
(2018)Tj
34.9018 0 Td
(2)Tj
0.7654 0 Td
(/)Tj
ET
endstream
endobj
34 0 obj
<>stream
endstream
endobj
35 0 obj
<>stream
endstream
endobj
36 0 obj
<>stream
endstream
endobj
37 0 obj
<>stream
endstream
endobj
38 0 obj
<>stream
endstream
endobj
39 0 obj
<>stream
endstream
endobj
40 0 obj
<>stream
endstream
endobj
41 0 obj
<>stream
endstream
endobj
42 0 obj
<>stream
endstream
endobj
43 0 obj
<>stream
endstream
endobj
44 0 obj
<>stream
endstream
endobj
45 0 obj
<>stream
endstream
endobj
46 0 obj
<>stream
endstream
endobj
47 0 obj
<>stream
endstream
endobj
48 0 obj
<>stream
endstream
endobj
49 0 obj
<>stream
endstream
endobj
50 0 obj
<>stream
endstream
endobj
51 0 obj
<>stream
endstream
endobj
52 0 obj
<>stream
BT
66.402 0 Td
(30)Tj
ET
Q
endstream
endobj
53 0 obj
<>/XObject<>>>/CropBox[0 0 612 792]/MediaBox[0 0 612 792]/Parent 8 0 R/Annots 55 0 R/Contents 56 0 R/TrimBox[0 0 612 792]>>
endobj
55 0 obj
[57 0 R 58 0 R 59 0 R 60 0 R 61 0 R]
endobj
57 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.g001)>>
endobj
58 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref026)>>
endobj
59 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.ref029)>>
endobj
60 0 obj
<>/Border[0 0 0]/Dest(Rpone.0193049.g001)>>
endobj
61 0 obj
<>/Border[0 0 0]/A 62 0 R>>
endobj
62 0 obj
<>
endobj
56 0 obj
[63 0 R 64 0 R 65 0 R 66 0 R 67 0 R]
endobj
63 0 obj
<>stream
q
0.83 0.64 0.02 0 k
360.6236 380.9197 m
387.0425 380.9197 l
h
f*
386.8157 367.937 m
396.2835 367.937 l
h
f*
401.4992 367.937 m
411.0236 367.937 l
h
f*
464.0882 302.9102 m
489.4299 302.9102 l
h
f*
0 g
1 j
1 J
0 w
10 0 0 10 200.0125 707.4141 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(inference)Tj
3.9344 0 Td
(from)Tj
2.2054 0 Td
(trajectory)Tj
4.0591 0 Td
(data.)Tj
2.143 0 Td
(As)Tj
1.2643 0 Td
(described)Tj
4.0535 0 Td
(in)Tj
1.0318 0 Td
(the)Tj
1.474 0 Td
(following)Tj
3.9741 0 Td
(section,)Tj
3.2769 0 Td
(these)Tj
2.2563 0 Td
(models)Tj
3.1011 0 Td
(are)Tj
1.4513 0 Td
(techni-)Tj
-34.2252 -1.3039 Td
(cally)Tj
2.0239 0 Td
(hybrid)Tj
2.8743 0 Td
(models)Tj
3.1067 0 Td
(that)Tj
1.7802 0 Td
(combine)Tj
3.7133 0 Td
(deterministic)Tj
5.5162 0 Td
(dynamics)Tj
4.0592 0 Td
(for)Tj
1.3833 0 Td
(animal)Tj
2.9707 0 Td
(motion)Tj
3.1691 0 Td
(with)Tj
1.9956 0 Td
(discrete)Tj
-32.5925 -1.2982 Td
(behavioral)Tj
4.3823 0 Td
(states;)Tj
2.6135 0 Td
(stochastic)Tj
4.1102 0 Td
(transitions)Tj
4.4731 0 Td
(between)Tj
3.5206 0 Td
(these)Tj
2.245 0 Td
(states)Tj
2.3868 0 Td
(are)Tj
1.4399 0 Td
(described)Tj
4.0422 0 Td
(in)Tj
1.0262 0 Td
(a)Tj
0.6463 0 Td
(Generalized)Tj
5.0059 0 Td
(Lin-)Tj
-35.892 -1.2983 Td
(ear)Tj
1.4456 0 Td
(Model)Tj
2.8176 0 Td
(framework.)Tj
-3.067 -1.2983 Td
(This)Tj
1.9899 0 Td
(paper)Tj
2.4831 0 Td
(is)Tj
0.8504 0 Td
(structured)Tj
4.3483 0 Td
(as)Tj
1.0148 0 Td
(follows.)Tj
3.2825 0 Td
(The)Tj
1.7858 0 Td
(next)Tj
1.9559 0 Td
(section)Tj
3.0444 0 Td
(introduces)Tj
4.4787 0 Td
(the)Tj
1.4684 0 Td
(rationale)Tj
3.753 0 Td
(behind)Tj
3.0217 0 Td
(the)Tj
-34.6731 -1.3039 Td
(modeling)Tj
4.0195 0 Td
(approach)Tj
3.9514 0 Td
(that)Tj
1.7745 0 Td
(we)Tj
1.3266 0 Td
(propose,)Tj
3.651 0 Td
(specifies)Tj
3.549 0 Td
(the)Tj
1.4683 0 Td
(model)Tj
2.7382 0 Td
(mathematically,)Tj
6.5707 0 Td
(and)Tj
1.7291 0 Td
(discusses)Tj
3.8325 0 Td
(the)Tj
1.4683 0 Td
(for-)Tj
-36.0791 -1.2982 Td
(ward)Tj
2.2336 0 Td
(\(simulation\))Tj
5.1817 0 Td
(and)Tj
1.7292 0 Td
(inverse)Tj
3.067 0 Td
(\(parameter)Tj
4.6545 0 Td
(inference\))Tj
4.2746 0 Td
(approaches.)Tj
4.9663 0 Td
(The)Tj
1.7801 0 Td
(second)Tj
3.0104 0 Td
(half)Tj
1.7178 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(paper)Tj
-35.0983 -1.2983 Td
(focuses)Tj
3.1124 0 Td
(on)Tj
1.2756 0 Td
(three)Tj
2.262 0 Td
(examples:)Tj
4.1385 0 Td
(two)Tj
1.7065 0 Td
(synthetic)Tj
3.8211 0 Td
(toy)Tj
1.4796 0 Td
(models)Tj
3.1068 0 Td
(of)Tj
1.0148 0 Td
(animal)Tj
2.9707 0 Td
(behavior)Tj
3.7133 0 Td
(\(ant)Tj
1.8369 0 Td
(motion,)Tj
3.3959 0 Td
(bacterial)Tj
-33.8341 -1.3039 Td
(chemotaxis\))Tj
5.0853 0 Td
(to)Tj
1.0261 0 Td
(showcase)Tj
3.9401 0 Td
(and)Tj
1.7292 0 Td
(validate)Tj
3.3051 0 Td
(the)Tj
1.474 0 Td
(inference)Tj
3.9345 0 Td
(as)Tj
1.0148 0 Td
(well)Tj
1.8198 0 Td
(as)Tj
1.0148 0 Td
(illustrate)Tj
3.6964 0 Td
(an)Tj
1.2019 0 Td
(interesting)Tj
4.49 0 Td
(extension)Tj
-33.732 -1.2983 Td
(to)Tj
1.0261 0 Td
(coarse-grained)Tj
6.1285 0 Td
(behavioral)Tj
4.388 0 Td
(states,)Tj
2.6248 0 Td
(and)Tj
1.7291 0 Td
(one)Tj
1.7008 0 Td
(real)Tj
1.6951 0 Td
(data)Tj
1.9106 0 Td
(example)Tj
3.5489 0 Td
(\(tracked)Tj
3.5376 0 Td
(zebrafish)Tj
3.8098 0 Td
(data\))Tj
2.2563 0 Td
(to)Tj
1.0262 0 Td
(illus-)Tj
-35.3818 -1.2983 Td
(trate)Tj
2.0466 0 Td
(inference)Tj
3.9344 0 Td
(without)Tj
3.3335 0 Td
(prior)Tj
2.2507 0 Td
(knowledge)Tj
4.5411 0 Td
(of)Tj
1.0148 0 Td
(laws)Tj
1.9502 0 Td
(of)Tj
1.0148 0 Td
(motion,)Tj
3.3902 0 Td
(as)Tj
1.0148 0 Td
(well)Tj
1.8199 0 Td
(as)Tj
1.0148 0 Td
(model)Tj
2.7439 0 Td
(selection)Tj
3.719 0 Td
(to)Tj
1.0261 0 Td
(iden-)Tj
-34.8148 -1.3039 Td
(tify)Tj
1.525 0 Td
(kinematic)Tj
4.2009 0 Td
(variables)Tj
3.719 0 Td
(that)Tj
1.7745 0 Td
(affect)Tj
2.3754 0 Td
(behavioral)Tj
4.3937 0 Td
(state)Tj
2.0353 0 Td
(switching.)Tj
ET
Q
q
1 j
1 J
0 w
11.9999 0 0 11.9999 200.0125 529.0015 cm
BT
/F2 1 Tf
1 TL
-0.0033 Tc
0 0 Td
(Methods)Tj
4.3086 0 Td
(and)Tj
1.989 0 Td
(models)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 200.0125 511.9937 cm
BT
/F6 1 Tf
1 TL
-0.004 Tc
0 0 Td
(The)Tj
1.7801 0 Td
(main)Tj
2.2904 0 Td
(goal)Tj
1.8708 0 Td
(of)Tj
1.0148 0 Td
(this)Tj
1.6781 0 Td
(work)Tj
2.2791 0 Td
(is)Tj
0.8447 0 Td
(to)Tj
1.0261 0 Td
(develop)Tj
3.3222 0 Td
(a)Tj
0.652 0 Td
(probabilistic)Tj
5.193 0 Td
(approach)Tj
3.9515 0 Td
(for)Tj
1.3889 0 Td
(understanding)Tj
6.0662 0 Td
(the)Tj
1.4683 0 Td
(behav-)Tj
-34.8262 -1.2982 Td
(ior)Tj
1.3662 0 Td
(and)Tj
1.7292 0 Td
(motion)Tj
3.1691 0 Td
(of)Tj
1.0148 0 Td
(animals,)Tj
3.5546 0 Td
(either)Tj
2.5285 0 Td
(in)Tj
1.0375 0 Td
(an)Tj
1.2018 0 Td
(individual)Tj
4.286 0 Td
(setting)Tj
2.8856 0 Td
(or)Tj
1.0999 0 Td
(as)Tj
1.0148 0 Td
(a)Tj
0.6519 0 Td
(group)Tj
2.6079 0 Td
(in)Tj
1.0375 0 Td
(which)Tj
2.6475 0 Td
(the)Tj
1.4684 0 Td
(animals)Tj
-33.3012 -1.3039 Td
(can)Tj
1.627 0 Td
(interact.)Tj
3.5093 0 Td
(We)Tj
1.6101 0 Td
(construe)Tj
3.6793 0 Td
(the)Tj
1.4683 0 Td
(behavior)Tj
4.4958 0 Td
(very)Tj
1.9275 0 Td
(broadly,)Tj
3.4923 0 Td
(as)Tj
1.0148 0 Td
(spanning)Tj
3.9118 0 Td
(the)Tj
1.4683 0 Td
(range)Tj
2.4605 0 Td
(from)Tj
2.2053 0 Td
(the)Tj
1.4683 0 Td
(occur-)Tj
-34.3386 -1.2983 Td
(rence)Tj
2.4094 0 Td
(of)Tj
1.0148 0 Td
(stereotyped)Tj
4.8245 0 Td
(events)Tj
2.7326 0 Td
(that)Tj
1.7745 0 Td
(can)Tj
1.6271 0 Td
(be)Tj
1.1508 0 Td
(identified)Tj
4.0592 0 Td
(by)Tj
1.1849 0 Td
(the)Tj
1.4683 0 Td
(body)Tj
2.2167 0 Td
(pose,)Tj
2.2507 0 Td
(specific)Tj
3.1918 0 Td
(interactions)Tj
4.9662 0 Td
(with)Tj
-34.8715 -1.2983 Td
(an)Tj
1.2018 0 Td
(environment,)Tj
5.6579 0 Td
(vocalization)Tj
5.0173 0 Td
(production,)Tj
4.9436 0 Td
(etc.,)Tj
1.8142 0 Td
(to)Tj
1.0261 0 Td
(internal)Tj
3.3562 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3981 0 Td
(that)Tj
1.7745 0 Td
(manifest)Tj
3.6623 0 Td
(with)Tj
-35.2457 -1.2982 Td
(the)Tj
1.4683 0 Td
(animal)Tj
2.9707 0 Td
(adopting)Tj
3.7814 0 Td
(a)Tj
0.6576 0 Td
(particular)Tj
4.1102 0 Td
(center-of-mass)Tj
6.1795 0 Td
(law)Tj
1.5817 0 Td
(of)Tj
1.0148 0 Td
(motion)Tj
3.1691 0 Td
(through)Tj
3.4469 0 Td
(space.)Tj
2.6022 0 Td
(We)Tj
1.6101 0 Td
(assume)Tj
3.1464 0 Td
(that)Tj
-35.7389 -1.3039 Td
(each)Tj
2.0295 0 Td
(animal)Tj
2.9651 0 Td
(switches)Tj
3.5659 0 Td
(between)Tj
3.532 0 Td
(the)Tj
1.4683 0 Td
(possible)Tj
3.4129 0 Td
(behaviors)Tj
4.0705 0 Td
(in)Tj
1.0375 0 Td
(a)Tj
0.652 0 Td
(stochastic)Tj
4.1215 0 Td
(fashion,)Tj
3.3789 0 Td
(with)Tj
1.9956 0 Td
(rates)Tj
2.1089 0 Td
(that)Tj
-34.3386 -1.2983 Td
(depend,)Tj
3.4128 0 Td
(in)Tj
1.0375 0 Td
(a)Tj
0.652 0 Td
(very)Tj
1.9219 0 Td
(general)Tj
3.1351 0 Td
(way)Tj
1.7914 0 Td
(that)Tj
1.7802 0 Td
(we)Tj
1.3266 0 Td
(seek)Tj
1.9219 0 Td
(to)Tj
1.0261 0 Td
(identify,)Tj
3.5206 0 Td
(on)Tj
1.2756 0 Td
(the)Tj
1.474 0 Td
(animal's)Tj
3.5433 0 Td
(current)Tj
3.1861 0 Td
(behavioral)Tj
4.388 0 Td
(state,)Tj
-35.3931 -1.2983 Td
(its)Tj
1.1451 0 Td
(location,)Tj
3.6794 0 Td
(and)Tj
1.7291 0 Td
(the)Tj
1.4683 0 Td
(location)Tj
3.4526 0 Td
(of)Tj
1.0148 0 Td
(other)Tj
2.3471 0 Td
(animals)Tj
3.3278 0 Td
(in)Tj
1.0375 0 Td
(a)Tj
0.652 0 Td
(group.)Tj
-18.6575 -1.3039 Td
(The)Tj
1.7858 0 Td
(expressive)Tj
4.2859 0 Td
(power)Tj
2.7269 0 Td
(of)Tj
1.0148 0 Td
(our)Tj
1.6215 0 Td
(model)Tj
2.7439 0 Td
(stems)Tj
2.4831 0 Td
(from)Tj
2.211 0 Td
(the)Tj
1.4684 0 Td
(combination)Tj
5.3461 0 Td
(of)Tj
1.0148 0 Td
(its)Tj
1.1451 0 Td
(stochastic)Tj
4.1216 0 Td
(and)Tj
1.7291 0 Td
(deter-)Tj
-34.8942 -1.2983 Td
(ministic)Tj
3.4695 0 Td
(components,)Tj
5.4028 0 Td
(as)Tj
1.0148 0 Td
(schematized)Tj
5.1364 0 Td
(in)Tj
0.83 0.64 0.02 0 k
1.0375 0 Td
(Fig)Tj
1.4796 0 Td
(1A)Tj
0 g
1.3777 0 Td
(and)Tj
1.7347 0 Td
(described)Tj
4.0479 0 Td
(in)Tj
1.0375 0 Td
(detail)Tj
2.4151 0 Td
(below.)Tj
2.8119 0 Td
(Formally,)Tj
-30.9654 -1.2982 Td
(our)Tj
1.6214 0 Td
(model)Tj
2.7382 0 Td
(belongs)Tj
3.2768 0 Td
(to)Tj
1.0205 0 Td
(a)Tj
0.652 0 Td
(class)Tj
2.0409 0 Td
(of)Tj
1.0148 0 Td
(hybrid)Tj
2.8743 0 Td
(models)Tj
3.1011 0 Td
([)Tj
0.83 0.64 0.02 0 k
(26)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(29)Tj
0 g
(])Tj
3.3165 0 Td
(with)Tj
1.9899 0 Td
(stochastic)Tj
4.1216 0 Td
(discrete)Tj
3.3108 0 Td
(states)Tj
2.3924 0 Td
(and)Tj
1.7292 0 Td
(deter-)Tj
-35.2004 -1.3039 Td
(ministic)Tj
3.4695 0 Td
(continuous)Tj
4.7395 0 Td
(dynamics,)Tj
4.2803 0 Td
(typically)Tj
3.5603 0 Td
(called)Tj
2.5228 0 Td
(Piecewise-Deterministic)Tj
9.9099 0 Td
(Markov)Tj
3.3732 0 Td
(Processes)Tj
-31.8555 -1.2983 Td
(\(PDMP\).)Tj
3.8721 0 Td
(While)Tj
2.6475 0 Td
(switching)Tj
4.0649 0 Td
(between)Tj
3.5262 0 Td
(behavioral)Tj
4.3937 0 Td
(states)Tj
2.3981 0 Td
(is)Tj
0.8447 0 Td
(stochastic,)Tj
4.3483 0 Td
(within)Tj
2.812 0 Td
(each)Tj
2.0296 0 Td
(behavioral)Tj
4.3937 0 Td
(state)Tj
-35.3308 -1.2983 Td
(the)Tj
1.4683 0 Td
(animal)Tj
2.9707 0 Td
(follows)Tj
3.0557 0 Td
(a)Tj
0.6576 0 Td
(certain)Tj
2.9877 0 Td
(law)Tj
1.5818 0 Td
(of)Tj
1.0148 0 Td
(motion)Tj
3.1691 0 Td
(that)Tj
1.7744 0 Td
(governs)Tj
3.3562 0 Td
(its)Tj
1.1452 0 Td
(kinematics)Tj
4.5581 0 Td
(\(e.g.,)Tj
2.126 0 Td
(its)Tj
1.1452 0 Td
(center-of-mass)Tj
-31.0108 -1.2982 Td
(trajectory\),)Tj
4.6204 0 Td
(as)Tj
1.0148 0 Td
(specified)Tj
3.7134 0 Td
(by)Tj
1.1792 0 Td
(a)Tj
0.6519 0 Td
(system)Tj
2.9367 0 Td
(of)Tj
1.0148 0 Td
(ordinary)Tj
3.6963 0 Td
(differential)Tj
4.6035 0 Td
(equations.)Tj
4.3143 0 Td
(This)Tj
1.9842 0 Td
(is)Tj
0.8504 0 Td
(a)Tj
0.6463 0 Td
(powerful)Tj
3.7871 0 Td
(gener-)Tj
-35.0133 -1.3039 Td
(alization)Tj
3.6339 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(traditional)Tj
4.422 0 Td
(models,)Tj
3.3278 0 Td
(schematized)Tj
5.1421 0 Td
(for)Tj
1.3833 0 Td
(comparison)Tj
4.9776 0 Td
(in)Tj
0.83 0.64 0.02 0 k
1.0374 0 Td
(Fig)Tj
1.4797 0 Td
(1B)Tj
0 g
(:)Tj
1.4967 0 Td
(in)Tj
1.0375 0 Td
(either)Tj
2.5341 0 Td
(zonal)Tj
2.3698 0 Td
(or)Tj
-35.3251 -1.2983 Td
(force)Tj
2.2336 0 Td
(field)Tj
1.9729 0 Td
(models,)Tj
3.3279 0 Td
(the)Tj
1.4683 0 Td
(focal)Tj
2.1203 0 Td
(animal)Tj
2.9707 0 Td
(always)Tj
2.8346 0 Td
(follows)Tj
3.0558 0 Td
(a)Tj
0.6576 0 Td
(single,)Tj
2.7553 0 Td
(universal)Tj
3.8551 0 Td
(law)Tj
1.5817 0 Td
(of)Tj
1.0148 0 Td
(motion)Tj
3.1691 0 Td
(that)Tj
1.7801 0 Td
(cannot)Tj
-34.7978 -1.2983 Td
(depend)Tj
3.1861 0 Td
(on)Tj
1.2756 0 Td
(the)Tj
1.4683 0 Td
(behavioral)Tj
4.3937 0 Td
(state.)Tj
2.262 0 Td
(Similarly,)Tj
4.0252 0 Td
(frameworks)Tj
4.9776 0 Td
(that)Tj
1.7801 0 Td
(model)Tj
2.7439 0 Td
(switching)Tj
4.0592 0 Td
(between)Tj
3.532 0 Td
(behav-)Tj
-33.7037 -1.3039 Td
(ioral)Tj
2.0466 0 Td
(states)Tj
2.3924 0 Td
(of)Tj
1.0148 0 Td
(individual)Tj
4.2916 0 Td
(animals)Tj
3.3279 0 Td
(often)Tj
2.2847 0 Td
(ignore)Tj
2.8006 0 Td
(the)Tj
1.474 0 Td
(fact)Tj
1.6611 0 Td
(that)Tj
1.7801 0 Td
(this)Tj
1.6725 0 Td
(behavior)Tj
3.7133 0 Td
(plays)Tj
2.2337 0 Td
(out)Tj
1.5534 0 Td
(in)Tj
1.0318 0 Td
(physical)Tj
-33.2785 -1.2983 Td
(space)Tj
2.3754 0 Td
(and)Tj
1.7348 0 Td
(is)Tj
0.8447 0 Td
(modulated)Tj
4.5354 0 Td
(by)Tj
1.1792 0 Td
(the)Tj
1.4683 0 Td
(ever-changing)Tj
5.9244 0 Td
(spatial)Tj
2.7779 0 Td
(relationship)Tj
4.989 0 Td
(between)Tj
3.5263 0 Td
(the)Tj
1.474 0 Td
(focal)Tj
2.1146 0 Td
(animal,)Tj
3.1975 0 Td
(its)Tj
-36.1415 -1.2982 Td
(conspecifics,)Tj
5.2553 0 Td
(and)Tj
1.7292 0 Td
(its)Tj
1.1452 0 Td
(environment.)Tj
5.6579 0 Td
(Our)Tj
1.8652 0 Td
(model)Tj
2.7439 0 Td
(brings)Tj
2.7439 0 Td
(together)Tj
3.532 0 Td
(the)Tj
1.4683 0 Td
(analysis)Tj
3.3278 0 Td
(of)Tj
1.0148 0 Td
(discrete)Tj
3.3166 0 Td
(behav-)Tj
-33.8001 -1.3039 Td
(ioral)Tj
2.0466 0 Td
(states)Tj
2.3924 0 Td
(and)Tj
1.7291 0 Td
(motion)Tj
3.1691 0 Td
(through)Tj
3.4469 0 Td
(space)Tj
2.3754 0 Td
(in)Tj
1.0375 0 Td
(a)Tj
0.6576 0 Td
(single)Tj
2.5285 0 Td
(probabilistic)Tj
5.1931 0 Td
(framework.)Tj
-23.38 -1.2983 Td
(Given)Tj
2.6305 0 Td
(our)Tj
1.6271 0 Td
(very)Tj
1.9219 0 Td
(general)Tj
3.135 0 Td
(notion)Tj
2.897 0 Td
(of)Tj
1.0148 0 Td
(what)Tj
2.1657 0 Td
(constitutes)Tj
4.5297 0 Td
(a)Tj
0.6577 0 Td
(behavior,)Tj
4.7225 0 Td
(there)Tj
2.262 0 Td
(exists)Tj
2.3981 0 Td
(no)Tj
1.2756 0 Td
(method)Tj
3.3221 0 Td
(for)Tj
-35.7559 -1.2983 Td
(automatically)Tj
5.6068 0 Td
(detecting)Tj
3.9005 0 Td
(and)Tj
1.7291 0 Td
(annotating)Tj
4.5694 0 Td
(such)Tj
2.058 0 Td
(behaviors)Tj
4.0762 0 Td
(in)Tj
1.0318 0 Td
(a)Tj
0.6576 0 Td
(way)Tj
1.7915 0 Td
(that)Tj
1.7801 0 Td
(is)Tj
0.8448 0 Td
(independent)Tj
5.278 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(ani-)Tj
-35.807 -1.3039 Td
(mal)Tj
1.7177 0 Td
(species,)Tj
3.2202 0 Td
(experimental)Tj
5.4595 0 Td
(setup,)Tj
2.5738 0 Td
(and)Tj
1.7291 0 Td
(the)Tj
1.474 0 Td
(biological)Tj
4.0876 0 Td
(question)Tj
3.6623 0 Td
(being)Tj
2.4265 0 Td
(asked.)Tj
2.6928 0 Td
(For)Tj
1.6215 0 Td
(example,)Tj
3.7757 0 Td
(when)Tj
-34.4407 -1.2983 Td
(describing)Tj
4.3823 0 Td
(ants,)Tj
2.0863 0 Td
(the)Tj
1.4683 0 Td
(most)Tj
2.2053 0 Td
(coarse-grained)Tj
6.1285 0 Td
(and)Tj
1.7291 0 Td
(easily)Tj
2.4095 0 Td
(observable)Tj
4.456 0 Td
(behaviors)Tj
4.0705 0 Td
(might)Tj
2.5965 0 Td
(be)Tj
1.1509 0 Td
(the)Tj
1.4683 0 Td
(modes)Tj
-34.1515 -1.2982 Td
(of)Tj
1.0147 0 Td
(locomotion)Tj
4.8473 0 Td
(\(such)Tj
2.4037 0 Td
(as)Tj
1.0148 0 Td
(stopping)Tj
3.702 0 Td
(and)Tj
1.7292 0 Td
(moving)Tj
3.2825 0 Td
(forward\);)Tj
3.9684 0 Td
(however,)Tj
3.8381 0 Td
(in)Tj
1.0375 0 Td
(other)Tj
2.3471 0 Td
(experimental)Tj
5.4538 0 Td
(setups)Tj
-34.6391 -1.2983 Td
(we)Tj
1.3266 0 Td
(might)Tj
2.5965 0 Td
(want)Tj
2.1883 0 Td
(to)Tj
1.0205 0 Td
(sidestep)Tj
3.4015 0 Td
(motion)Tj
3.1691 0 Td
(details)Tj
2.778 0 Td
(and)Tj
1.7291 0 Td
(instead)Tj
3.0784 0 Td
(focus)Tj
2.3301 0 Td
(on)Tj
1.2756 0 Td
(anntenation)Tj
5.0739 0 Td
(and)Tj
1.7292 0 Td
(grooming)Tj
-31.6968 -1.3039 Td
(behaviors)Tj
4.0705 0 Td
(triggered)Tj
3.8324 0 Td
(by)Tj
1.1849 0 Td
(exposure)Tj
3.8154 0 Td
(to)Tj
1.0261 0 Td
(specific)Tj
3.1918 0 Td
(pathogens.)Tj
4.5241 0 Td
(In)Tj
1.1111 0 Td
(the)Tj
1.4684 0 Td
(absence)Tj
3.3392 0 Td
(of)Tj
1.0148 0 Td
(generic)Tj
3.1464 0 Td
(algorithms)Tj
4.5184 0 Td
(for)Tj
-36.2435 -1.2983 Td
(behavioral)Tj
4.3936 0 Td
(annotation,)Tj
4.8359 0 Td
(we)Tj
1.3266 0 Td
(start)Tj
1.9842 0 Td
(here)Tj
1.9616 0 Td
(by)Tj
1.1849 0 Td
(assuming)Tj
4.0025 0 Td
(that)Tj
1.7801 0 Td
(we)Tj
1.3266 0 Td
(are)Tj
1.4514 0 Td
(already)Tj
3.1067 0 Td
(given)Tj
2.3811 0 Td
(individual)Tj
4.286 0 Td
(animal)Tj
-34.0212 -1.2982 Td
(motion)Tj
3.1634 0 Td
(trajectories)Tj
4.6601 0 Td
(that)Tj
1.7745 0 Td
(have)Tj
2.0636 0 Td
(been)Tj
2.1203 0 Td
(annotated)Tj
4.2406 0 Td
(with)Tj
1.9956 0 Td
(behaviors)Tj
4.0762 0 Td
(of)Tj
1.0148 0 Td
(interest;)Tj
3.4356 0 Td
(this)Tj
1.6724 0 Td
(input)Tj
2.3811 0 Td
(data)Tj
1.9162 0 Td
(is)Tj
-34.5144 -1.3039 Td
(obtained)Tj
3.7303 0 Td
(from)Tj
2.2054 0 Td
(raw)Tj
1.7121 0 Td
(observations)Tj
5.261 0 Td
(in)Tj
1.0318 0 Td
(a)Tj
0.6577 0 Td
(problem-specific)Tj
6.9335 0 Td
(way)Tj
1.7915 0 Td
(that)Tj
1.7801 0 Td
(we)Tj
1.3266 0 Td
(do)Tj
1.2529 0 Td
(not)Tj
1.5761 0 Td
(address)Tj
3.2258 0 Td
(here.)Tj
2.1883 0 Td
(Start-)Tj
-34.6731 -1.2983 Td
(ing)Tj
1.5023 0 Td
(with)Tj
1.9956 0 Td
(such)Tj
2.0579 0 Td
(input)Tj
2.3811 0 Td
(data,)Tj
2.1373 0 Td
(our)Tj
1.6271 0 Td
(framework)Tj
4.6148 0 Td
(will)Tj
1.6667 0 Td
(then)Tj
2.0183 0 Td
(proceed)Tj
3.4129 0 Td
(to)Tj
1.0261 0 Td
(identify)Tj
3.2938 0 Td
(how)Tj
1.9389 0 Td
(the)Tj
1.4684 0 Td
(animals)Tj
3.3278 0 Td
(switch)Tj
ET
Q
q
1 j
1 J
0 w
576 737.1 m
36 737.1 l
36 737.6 l
576 737.6 l
f*
36 741.2598 107.1496 23.6976 re
W* n
q
107.0929 0 0 23.6409 36 741.3165 cm
q
/I0 Do
Q
Q
Q
q
0 0 612 792 re
W* n
1 j
1 J
0 w
7.9999 0 0 7.9999 441.5811 745.7952 cm
BT
/F0 1 Tf
1 TL
-0.005 Tc
0 0 Td
(Probabilist)Tj
4.5709 0 Td
(ic)Tj
0.9283 0 Td
(models)Tj
3.3874 0 Td
(of)Tj
1.0418 0 Td
(animal)Tj
3.111 0 Td
(behavio)Tj
3.43 0 Td
(r)Tj
ET
Q
q
1 j
1 J
0 w
36 48.0002 m
576 48.0002 l
576 47.5002 l
36 47.5002 l
f*
0.83 0.64 0.02 0 k
81.5811 34.9228 m
237.8835 34.9228 l
h
f*
0 g
7.9999 0 0 7.9999 36 36 cm
BT
/F0 1 Tf
1 TL
-0.005 Tc
0 0 Td
(PLOS)Tj
2.8559 0 Td
(ONE)Tj
2.3669 0 Td
(|)Tj
0.83 0.64 0.02 0 k
0.4748 0 Td
(https://doi.or)Tj
5.4142 0 Td
(g/10.137)Tj
3.8198 0 Td
(1/journal.po)Tj
5.1236 0 Td
(ne.01930)Tj
4.089 0 Td
(49)Tj
0 g
2.0906 0 Td
(March)Tj
2.9622 0 Td
(7,)Tj
1.0418 0 Td
(2018)Tj
34.9018 0 Td
(3)Tj
0.7654 0 Td
(/)Tj
ET
endstream
endobj
64 0 obj
<>stream
endstream
endobj
65 0 obj
<>stream
endstream
endobj
66 0 obj
<>stream
endstream
endobj
67 0 obj
<>stream
BT
66.402 0 Td
(30)Tj
ET
Q
endstream
endobj
68 0 obj
<>/XObject<>>>/CropBox[0 0 612 792]/MediaBox[0 0 612 792]/Parent 8 0 R/Annots 72 0 R/Contents 73 0 R/TrimBox[0 0 612 792]>>
endobj
71 0 obj
<>stream
x\GƿXc/&jEQDw{b7]XQ(
RE=q{AT02w̙3ݻg̙ ~8;)NjRuʗ3M$@$@$@$@$@!2]#YIeQ:rH,L %XoVtt+WΝ?ҥGż}+
`77W{zz;H< ;\WGNppcG/\ !O?~(LSY$@$@$@$@$@$]x]xxÇaԜ:u̙3]i5EDDݻw.(Y>ް^~#l%A 9!!w9}ѣG-/_B+Q /# [a`|3~y2e47=y``BQEHYlR˗766Y̐hݺE>刊"CnݪDJff֥S[2hQ ٴy'ƍ/Q"ET2hyF[[5jVkm]uܡI%pԭ[v۶mIoݲƵ/_PH*y=z.\T
4XsݡIHHHHK`aSCUk_|Yȁ)ereˊӥkԠ*bRFX4&HHHHHH_!qIQ#÷ldRJt&}ڷ?⢬5z\r sR-g
m׃QQQBǏmڨQڴiɿm۲ɈD9i ۹mkiҤV2XZX۳')B^EF_B2 {&T*\oӳgySOs*Xpɚ):5T!1A$@$@$@$@$@
tADwקO,Ys
Y<(HQ'˛WDԩseTҠΜ<>BDb\rҤIϒ9M7qޤU%1A$@$@$@$@$@ -Kff,lkW';DZ$?wwP>9sdSr`W.YQQX֯wy(ӥM[r#c֥tL $ [Fo^{w9==aeΔ~O<*vtSX1Qk7z@hs K
E߅أu(/r &m=EFDZx ˔1AuF"ݻS'WdH0yRYЯ3YҰSjת%gʔik,i f`ݼqm_U12kJٛ7oXJ6DDT*W2ܵc;BV9}d&3g~?s(YYqqMCLi &bwãM˖ӿʔٺe˫D4NC&+W,ϖ5l
Ȭ^1wB%TE4<ۧ,F/]h*~_ f91tNtUِ ~qΜŋbJڊ/nbl\zff&&؟q5j/s}!0>bcK}"6D$@$@$@$@$@Y洤d3g=