132:
localization and a wider range of sensitivity to sound. The movement of the wings during flight also plays a role, since sound thresholds change with wing position. The neural mechanisms for triggering the acoustic startle response are partially understood. However, there is little known about the motor control of flight that ultrasound initiates.
151:
245:
In praying mantises, ultrasound avoidance behaviors are non-directional turns or power dives that are very effective in preventing capture by bats. The mantis ear, located in the midline between the metathoracic (third) legs, comprises two tympana within an auditory chamber that enhances sensitivity.
163:
Crickets are preyed on by bats during the night while they fly from one place to another. Avoidance behaviors by crickets were first reported in 1977 by A. V. Popov and V. F. Shuvalov. They also demonstrated that crickets, like moths, fly away from bats once they've heard their echolocating calls, an
135:
Further research has shown that many species of moths are sensitive to ultrasound. Sensitivities for ultrasound change according to the environment the moth thrives in, and the moth can even change its own sensitivity if it is preyed upon by bats with different echolocating calls. Such is the case of
246:
A bilaterally symmetrical pair of auditory interneurons, 501-T3, accurately track the ultrasonic calls during the early stages of a bat attack. Because 501-T3 stops firing just before the evasive response starts, it may be involved in triggering the behavior. The praying mantis ear first appeared
206:
of sorts; in the cricket, int-1 is a bat detector when the cricket is in flight and the interneuron's activity reaches a specific threshold. If these conditions are met, the magnitude of the sound is linearly proportional to the magnitude of the avoidance response. This research also demonstrated
175:
fashion. This is understandable since crickets don't only need to listen to bats, but also to each other. Crickets have broad frequency sensitivity to different types of echolocating calls. One specific auditory interneuron, the AN2 interneuron, exhibits remarkably rapid responses to echolocating
131:
The moth's body axis allows it to be more sensitive to sounds coming from particular directions. Their ears, on either side of the metathorax, have two sensory cells within the membranes. Though the tuning curves of these cells are identical, the sensitivity thresholds differ, allowing for sound
168:. The cricket will steer itself away from the source of the sound within a very short time frame (40β80 ms). The response is evoked by brief ultrasonic pulses in the 20 to 100 kHz range, pulses which fall within the range of bat ultrasonic echolocating calls (20β100 kHz).
253:
Arctiid moths use a very different, but highly effective defense against bats. They produce loud ultrasonic clicks in response to ultrasound. Depending on the species of moth and its ecology, the clicks may work by startling the bat, by jamming its echolocation system, or by warning of
193:
cricket, two ascending interneurons carry information to the brain - one carries information about cricket song (around 5 kHz) while the other gets excited at ultrasound and other high frequencies (15β100 kHz). The ultrasound-sensitive interneuron - labeled
115:
It was found that the moths' responses vary according to ultrasound intensity, diving towards the ground if the pulse was of a high amplitude, or flying directly away from the sound source if the sound amplitude was low (if the sound was softer). Acoustic
265:) have sensitive ears on their wings. Ultrasound causes flying lacewings to fold their wings and drop, an effective maneuver for evading capture by bats. Some tettigoniids use a similar strategy, although other species respond much like crickets.
92:
made the association between the moth's high-pitched sounds and the high-pitched bat calls and wondered whether the moths would be able to hear it. However, it was not until the early 1960s that
Kenneth Roeder et al. made the first
112:; they fly away from the source of the sound and will only have the diving behavior considered above when the sound is too loudβor when, in a natural setting, the bat would be presumably too close to simply fly away.
201:
Stimulating int-1 by current injection is sufficient to initiate negative phonotaxis, while hyperpolarizing int-1 effectively cancels the turning response to ultrasound. Due to this, int-1 has been proposed to be a
423:
268:
Several other insects have sensitive ultrasonic hearing that probably is used in bat evasion, but direct evidence is not yet available. These include scarab beetles, tiger beetles and a parasitoid fly (
222:. Furthermore, it has been found that the ultrasound avoidance response is restricted to when the crickets are in flight: that is, the response is extinguished when the crickets are on the ground.
104:
Later research showed that moths responded to ultrasound with evasive movements. Moths, as do crickets and most insects that display bat avoidance behaviors, have tympanic organs that display
561:
Fullard, J. H.; Jackson, M. E.; Jacobs, D. S.; Pavey, C. R. & Burwell, C. J. (2008). "Surviving cave bats: auditory and behavioural defences in the
Australian noctuid moth,
225:
It has also been shown that short-winged crickets are less sensitive to ultrasound, but not to low frequencies, than their long-winged counterparts in a wing-dimorphic cricket,
237:(JH), is believed to play a role in whether the individual develops shorter or longer wings: if the individual has a higher level of JH, its wings will be shorter.
685:
Fullard, J. H.; Ratcliffe, J. M. & Guignion, C. (2005). "Sensory ecology of predatorβprey interactions: responses of the AN2 interneuron in the field cricket,
250:
120 million years ago, predating the appearance of echolocating bats by c. 50 million years, so its original function must be different from its current one.
778:
Hofstede, H. M.; Killow, J. & Fullard, J. H. (2009). "Gleaning bat echolocation calls do not elicit antipredator behaviour in the
Pacific field cricket,
1257:
Robert, D. & Hoy, R. R. (1998). "The evolutionary innovation of tympanal hearing in
Diptera". In: Hoy, R. R.; Popper, A. N. & Fay, R. R., editors.
1080:
Yager, D. D. & Svenson, G. J. (2008). "A phylogeny of mantis auditory systems based on morphological, molecular, physiological, and behavioral data".
37:
Although ultrasonic signals are used for echolocation by toothed whales, no known examples of ultrasonic avoidance in their prey have been found to date.
1270:
Boyan, G. S. & Miller, L. A. (1991). "Parallel processing of afferent input by identified interneurones in the auditory pathway of the noctuid moth
1005:
Triblehorn, J. D. & Yager, D. D. (2002). "Implanted electrode recordings from a praying mantis auditory interneuron during flying bat attacks".
88:
The idea that moths were able to hear the cries of echolocating bats dates back to the late 19th century. F. Buchanan White, in an 1877 letter to
34:
tuned to sense the bat's echolocating calls. The ultrasonic hearing is coupled to a motor response that causes evasion of the bat during flight.
1106:
Miller, L. A. (1984). "Hearing in green lacewings and their responses to the cries of bats". In: Canard, M.; SΓ©mΓ©ria, Y.; New, T. R., editors.
825:
Narbonne, R. & Pollack, G. S. (2008). "Developmental control of ultrasound sensitivity by a juvenile hormone analog in crickets (
1325:
216:
cricket, its broad sensitivity can be circumvented by the use of frequency-mismatched calls by part of bats like the gleaning bat,
207:
that the brain is necessary for the response, since decapitated crickets will fly, but show no avoidance response behaviors.
124:
located in a chamber formed by the wall of the abdomen and the tympanic membrane, are most sensitive to lower frequencies of
142:, which adapts its acoustic sensitivity according to the characteristics of the call of the bat inside the cave with them.
1320:
332:"Intense ultrasonic clicks from echolocating toothed whales do not elicit anti-predator responses or debilitate the squid
1093:
Conner, W. E. & Corcoran, A. J. (2012). "Sound strategies: the 65-million-year-old battle between bats and insects".
44:
era, (about 50 million years ago); antibat tactics should have evolved then. Antibat tactics are known in four orders of
171:
As opposed to moths, the cricket ear, located in the foreleg, is complex - having 70 receptors that are arranged in a
26:
predators. Ultrasound avoidance is known for several groups of insects that have independently evolved mechanisms for
735:
Nolen, T. G. & Hoy, R. R. (1984). "Initiation of behavior by single neurons: the role of behavioral context".
408:
391:
1119:
Libersat, F. & Hoy, R. R. (1991). "Ultrasonic startle behavior in bushcrickets
Orthoptera; Tettigoniidae".
1042:"Timing of praying mantis evasive responses during simulated bat attack sequences. When does the mantis dive?"
435:
Payne, R. S.; Roeder, K. D. & Wallman, J. (1966). "Directional
Sensitivity of the Ears of Noctuid Moths".
198:- has been demonstrated as both necessary and sufficient for negative phonotaxis by Nolen and Hoy in 1984:
80:). There are hypotheses of ultrasound avoidance being present in Diptera (flies) and Coleoptera (beetles).
212:
184:
643:
Boyan, G. S. & Fullard, J. H. (1986). "Interneurones responding to sound in the tobacco budworm moth
392:"How Some Insects Detect and Avoid Being Eaten by Bats: Tactics and Countertactics of Prey and Predator"
218:
744:
479:
138:
94:
40:
Ultrasonic hearing has evolved multiple times in insects: a total of 19 times. Bats appeared in the
1330:
165:
156:
109:
23:
1299:
987:
807:
714:
664:
625:
590:
27:
1291:
1240:
1195:
1154:
1063:
1022:
979:
940:
891:
846:
799:
760:
706:
582:
543:
495:
452:
365:
312:
117:
31:
183:
on a far lower number of interneurons that relay the receptors' information to the cricket's
22:
is an escape or avoidance reflex displayed by certain animal species that are preyed upon by
1283:
1230:
1185:
1146:
1053:
1014:
971:
930:
881:
838:
791:
752:
698:
656:
617:
574:
533:
487:
444:
403:
355:
347:
304:
234:
121:
57:
958:
Yager, D. D. & Hoy, R. R. (1987). "The midline metathoracic ear of the praying mantis,
195:
748:
483:
1133:
Schulze, W. & Schul, J. (2001). "Ultrasound avoidance behavior in the bush cricket
360:
331:
291:
Schulze, W. & Schul, J. (2001). "Ultrasound avoidance behaviour in the bushcricket
203:
98:
73:
1314:
30:. Insects have evolved a variety of ultrasound-sensitive ears based upon a vibrating
1303:
991:
811:
718:
668:
629:
842:
189:
594:
262:
255:
105:
69:
53:
909:
Triblehorn, J. D.; Ghose, K.; Bohn, K.; Moss, C. M. & Yager, D. D. (2008).
608:
Popov, A. V. & Shuvalov, V. F. (1977). "Phonotactic behavior of crickets".
795:
702:
125:
77:
61:
866:"Ultrasound-triggered, flight-gated evasive maneuvers in the praying mantis,
1190:
1173:
756:
172:
1158:
1067:
1026:
944:
850:
803:
710:
586:
491:
470:
Roeder, K. D. (1975). "Neural factors and evitability in insect behavior".
369:
351:
316:
1295:
1244:
1235:
1214:
1199:
1150:
1018:
983:
895:
764:
547:
538:
521:
499:
456:
308:
150:
886:
865:
65:
448:
1287:
975:
935:
910:
660:
621:
578:
230:
180:
45:
1058:
1041:
41:
522:"The Neuroethology of Acoustic Startle and Escape in Flying insects"
330:
Wilson, M.; Hanlon, R. T.; Tyack, P. L. & Madsen, P. T. (2007).
270:
149:
49:
647:(Noctuidae): morphological and physiological characteristics".
409:
10.1641/0006-3568(2001)051[0570:HSIDAA]2.0.CO;2
210:
Bats may have found ways to get around this system. In the
1174:"Ultrasound acoustic startle responses in scarab beetles"
1215:"Behavioral response to ultrasound in the tiger beetle,
1172:
Forrest, T. G.; Farris, H. E. & Hoy, R. R. (1995).
16:
Animal reflex to escape or avoid echolocating predators
1219:
Dow combines aerodynamic changes and sound production"
422:
White's reference can be found in the following link:
1261:. Heidelberg and New York: Springer-Verlag. p 197-227
864:
Yager, D. D.; May, M. L. & Fenton, M. B. (1990).
425:. His question is close to the ending of the letter.
911:"Free-flight encounters between the praying mantis
520:Hoy, R.; Nolen, T. & Brodfuehrer, P. (1989).
154:An adult male and a juvenile male of the species
689:to the echolocation calls of sympatric bats".
1110:. The Hague: Dr W. Junk Publishers, p 134-149
1040:Triblehorn, J. D. & Yager, D. D. (2005).
8:
1213:Yager, D. D. & Spangler, H. G. (1997).
680:
678:
1234:
1189:
1082:Biological Journal of the Linnean Society
1057:
934:
885:
537:
407:
390:Miller, L. A. & Surlykke, A. (2001).
359:
101:and were able to confirm this suspicion.
730:
728:
385:
383:
381:
379:
283:
515:
513:
511:
509:
241:Ultrasound avoidance in other insects
7:
1276:Journal of Comparative Physiology A
1223:The Journal of Experimental Biology
1178:The Journal of Experimental Biology
1139:The Journal of Experimental Biology
1121:Journal of Comparative Physiology A
1046:The Journal of Experimental Biology
1007:The Journal of Experimental Biology
923:The Journal of Experimental Biology
874:The Journal of Experimental Biology
784:Journal of Comparative Physiology A
691:Journal of Comparative Physiology A
567:The Journal of Experimental Biology
526:The Journal of Experimental Biology
472:The Journal of Experimental Biology
437:The Journal of Experimental Biology
14:
649:Journal of Comparative Physiology
610:Journal of Comparative Physiology
146:Ultrasound avoidance in crickets
297:Journal of Experimental Biology
128:(between 20 and 30 kHz.).
97:recordings of a noctuid moth's
843:10.1016/j.jinsphys.2008.09.004
295:(Orthoptera: Tettigoniidae)".
1:
247:
136:the Australian noctuid moth,
84:Ultrasound avoidance in moths
1259:Comparative Hearing: Insects
831:Journal of Insect Physiology
1095:Annual Review of Entomology
1347:
782:(Orthoptera: Gryllidae)".
870:(Gerst.). I. Free flight"
796:10.1007/s00359-009-0454-3
703:10.1007/s00359-005-0610-3
1326:Antipredator adaptations
964:Cell and Tissue Research
1191:10.1242/jeb.198.12.2593
913:Parasphendale agrionina
868:Parasphendale agrionina
757:10.1126/science.6505681
1135:Tettigonia viridissima
1108:Biology of Chrysopidae
827:Teleogryllus oceanicus
780:Teleogryllus oceanicus
687:Teleogryllus oceanicus
492:10.1002/jez.1401940106
352:10.1098/rsbl.2007.0005
293:Tettigonia viridissima
213:Teleogryllus oceanicus
185:central nervous system
160:
1236:10.1242/jeb.200.3.649
1151:10.1242/jeb.204.4.733
1019:10.1242/jeb.205.3.307
539:10.1242/jeb.146.1.287
309:10.1242/jeb.204.4.733
219:Nyctophilus geoffroyi
153:
120:in noctuid moths are
1321:Animal communication
1184:(Pt 12): 2593β2598.
1052:(Pt 10): 1867β1876.
887:10.1242/jeb.152.1.17
563:Speiredonia spectans
179:All these receptors
139:Speiredonia spectans
95:electrophysiological
20:Ultrasound avoidance
749:1984Sci...226..992N
645:Heliothis virescens
484:1975JEZ...194...75R
449:10.1242/jeb.44.1.17
166:negative phonotaxis
157:Gryllus bimaculatus
110:directional hearing
1288:10.1007/bf00224361
976:10.1007/bf00218944
936:10.1242/jeb.005736
661:10.1007/BF00603623
622:10.1007/BF00655876
579:10.1242/jeb.023978
573:(Pt 24): 3808β15.
161:
28:ultrasonic hearing
1229:(Pt 3): 649β659.
1217:Cicindela marutha
1059:10.1242/jeb.01565
1013:(Pt 3): 307β320.
929:(Pt 4): 555β562.
743:(4677): 992β994.
261:Green lacewings (
254:distastefulness (
118:sensory receptors
32:tympanic membrane
1338:
1307:
1262:
1255:
1249:
1248:
1238:
1210:
1204:
1203:
1193:
1169:
1163:
1162:
1130:
1124:
1117:
1111:
1104:
1098:
1091:
1085:
1078:
1072:
1071:
1061:
1037:
1031:
1030:
1002:
996:
995:
960:Mantis religiosa
955:
949:
948:
938:
917:Eptesicus fuscus
906:
900:
899:
889:
861:
855:
854:
822:
816:
815:
775:
769:
768:
732:
723:
722:
682:
673:
672:
640:
634:
633:
605:
599:
598:
558:
552:
551:
541:
517:
504:
503:
467:
461:
460:
432:
426:
420:
414:
413:
411:
387:
374:
373:
363:
327:
321:
320:
288:
249:
235:juvenile hormone
227:Grillus texensis
122:mechanoreceptors
1346:
1345:
1341:
1340:
1339:
1337:
1336:
1335:
1311:
1310:
1269:
1266:
1265:
1256:
1252:
1212:
1211:
1207:
1171:
1170:
1166:
1132:
1131:
1127:
1123:. 169: 507-514.
1118:
1114:
1105:
1101:
1092:
1088:
1079:
1075:
1039:
1038:
1034:
1004:
1003:
999:
957:
956:
952:
908:
907:
903:
863:
862:
858:
824:
823:
819:
777:
776:
772:
734:
733:
726:
684:
683:
676:
642:
641:
637:
607:
606:
602:
560:
559:
555:
519:
518:
507:
469:
468:
464:
434:
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429:
421:
417:
389:
388:
377:
340:Biology Letters
329:
328:
324:
290:
289:
285:
280:
243:
148:
86:
74:green lacewings
17:
12:
11:
5:
1344:
1342:
1334:
1333:
1328:
1323:
1313:
1312:
1309:
1308:
1272:Noctua pronuba
1264:
1263:
1250:
1205:
1164:
1125:
1112:
1099:
1086:
1084:. 94: 541-568.
1073:
1032:
997:
970:(3): 531β541.
950:
901:
856:
837:(12): 1552β6.
817:
770:
724:
674:
655:(3): 391β404.
635:
600:
553:
505:
462:
427:
415:
402:(7): 570β581.
375:
346:(3): 225β227.
334:Loligo pealeii
322:
303:(4): 733β740.
282:
281:
279:
276:
242:
239:
204:command neuron
176:call stimuli.
147:
144:
99:auditory nerve
85:
82:
15:
13:
10:
9:
6:
4:
3:
2:
1343:
1332:
1329:
1327:
1324:
1322:
1319:
1318:
1316:
1305:
1301:
1297:
1293:
1289:
1285:
1282:(6): 727β38.
1281:
1277:
1273:
1268:
1267:
1260:
1254:
1251:
1246:
1242:
1237:
1232:
1228:
1224:
1220:
1218:
1209:
1206:
1201:
1197:
1192:
1187:
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1179:
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1136:
1129:
1126:
1122:
1116:
1113:
1109:
1103:
1100:
1096:
1090:
1087:
1083:
1077:
1074:
1069:
1065:
1060:
1055:
1051:
1047:
1043:
1036:
1033:
1028:
1024:
1020:
1016:
1012:
1008:
1001:
998:
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989:
985:
981:
977:
973:
969:
965:
961:
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951:
946:
942:
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932:
928:
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918:
914:
905:
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888:
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869:
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848:
844:
840:
836:
832:
828:
821:
818:
813:
809:
805:
801:
797:
793:
790:(8): 769β76.
789:
785:
781:
774:
771:
766:
762:
758:
754:
750:
746:
742:
738:
731:
729:
725:
720:
716:
712:
708:
704:
700:
697:(7): 605β18.
696:
692:
688:
681:
679:
675:
670:
666:
662:
658:
654:
650:
646:
639:
636:
631:
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623:
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584:
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568:
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557:
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531:
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523:
516:
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477:
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405:
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397:
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96:
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83:
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67:
63:
59:
55:
51:
47:
43:
38:
35:
33:
29:
25:
21:
1279:
1275:
1271:
1258:
1253:
1226:
1222:
1216:
1208:
1181:
1177:
1167:
1142:
1138:
1134:
1128:
1120:
1115:
1107:
1102:
1094:
1089:
1081:
1076:
1049:
1045:
1035:
1010:
1006:
1000:
967:
963:
959:
953:
926:
922:
916:
915:and the bat
912:
904:
877:
873:
867:
859:
834:
830:
826:
820:
787:
783:
779:
773:
740:
736:
694:
690:
686:
652:
648:
644:
638:
613:
609:
603:
570:
566:
562:
556:
529:
525:
478:(1): 75β88.
475:
471:
465:
443:(1): 17β31.
440:
436:
430:
418:
399:
395:
343:
339:
333:
325:
300:
296:
292:
286:
269:
267:
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252:
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226:
224:
217:
211:
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