325:
beta pathway, the signal can then pass a chemical synapse, the information is sent to a sensory interneuron (SI) of either type A (which fires phasically in response to input) or type C, (which fire tonically). From there, the signal is sent across an electrical synapse to the LGI. The LGI can also receive the sensory input directly through an electrical synapse by way of the alpha pathway. This bypasses the SIs. The LGI can pass the signal to the MoG's by two paths one of which passes the signal to the Motor Giant through direct electrical synapses. The MoG then passes the signal to the FF muscles. It can also use a second route across a fast electrical synapse to a premotor interneuron called the segmental giant (SG), two of which are located in each segment. The SGs then relays the signal to fast flexor motor neurons. The end result is a powerful and rapid flexion.
309:(also called septate synapses). As a result, the LGI functions as one giant, continuous neuron, such as the MG. The LGI's large diameter and the efficiency and speed of electrical synapses, make the LGI-mediated escape especially rapid and effective. The LGI only innervates the first three rostral segments of the tail and is activated within 10 ms from when a mechanical stimulus is presented to the abdomen. When flexion occurs, only the first three segments flex and the tail fan is not directed under the body and forward but rather straight down. The unflexed segments increase the length of the paddle. The result is the rear end of the animal is directed upward and forward causing the animal to tumble or somersault forward. The tail is then rapidly extended, and this is usually followed by directed swimming.
275:. These fibers terminate in the last abdominal segment. The response is triggered by abrupt tactile stimuli to the head or alarming visual stimuli. This results in the firing of all motor giant (MoG) neurons and the flexion of all the phasic fast flexor (FF) muscles in the abdomen, which rapidly curls the tail fan and last three segments underneath the crayfish. This produces a thrust that directs water towards the offending stimulus while propelling the crayfish directly backward. This behavior is then followed by the extension of the tail, which prepares it for swimming or the execution of another tail-flip. The extension and swimming phases will be discussed in the next escape variant.
391:
the inhibitory effect is lost and the LGI in severed abdomen generates strong spikes since its threshold had decreased substantially. This is an indication that the behavior is mediated in the brain or the thorax of the crayfish. Most non-giant and MGI tail flips are also suppressed by restraint, so only a few non-giant systems are active to allow the crayfish to carefully wriggle free. The setup of this system seems to be designed to prevent the unnecessary use of all or nothing tail flips or ineffective tail flips when the situation requires more careful maneuvering.
148:. The paper stated that command neurons were neurons (or small sets of neurons) carrying the entire command signal for a natural behavioral response. According to the authors, command neurons were both necessary and sufficient in the production of a behavioral response. The concept of command neuron-mediated behaviors was both ground breaking and controversial, since determining command neuron-mediated behaviors was a problematic process due to difficulties in understanding and interpreting anatomical and behavioral data.
201:
pairs of walking legs. This is the crayfish's primary mode of locomotion. The abdominal section of the crayfish is divided into seven segments. These segments are flexibly interconnected, forming the tail. Normally, the tail is held in an extended position to aid in maneuvering and balancing. The first five segments are similar and the two terminal segments are modified into a tail fan, a region with high surface area that acts as the blade of a paddle in the escape response. This region contains the
288:
interneurons, most likely because they do not produce depolarizations in the sensory neurons that are above the thresholds required to initiate these behaviors. Instead they involve one or a few of the smaller neurons innervating the tail. The latency periods for these escape mechanisms are longer, ranging from 80 to 500 ms. However, this slower swimming behavior allows for flexibility, since the crayfish can use visual stimuli and steer itself in a selected direction.
98:
1586:
418:
decreased. Aggressive dominant males have a moderate reduction in tail flips while the subordinates have a much lower occurrence of tail flips. This presents a paradox since this means that subordinates are more likely to get killed. However, it was found that subordinates are more likely to use non-giant-mediated escape, indicating that the reduction in tail flips and enhancement of non giant escape is adaptive.
333:
prevent these events. The extensor muscles are inhibited to prevent competition between the extensor and flexors. Rapid IPSPs presented at the muscle receptor organ (MRO) prevent the stretch receptor from initiating extension while they are also presented at the fast extensor motor neurons and the fast extensor muscles. The circuits responsible for slow flexion and extension are also inhibited.
20:
1598:
360:
after the completion of the tail flip. The non-giant swimming occurs independently of the LGI response since direct stimulation of the LGI with the electrodes results in a tail flip but not the subsequent non-giant swimming. This swimming response seems to be a fixed action pattern mediated by a central pattern generator since it does not require
422:
to be mediated by different cell receptors. Differences in the effects of serotonin on the behavior of the crayfish seem to be the result of differences in the populations of these receptors. It is unknown how these modulation processes convey the information to the LG, and how the behavioral changes are precipitated.
296:
by non-giant circuitry have lower amplitudes and longer durations while the recordings in the FF muscles are more erratic and have smaller amplitudes since they receive smaller EPSPs from the MoGs. So while it is hard to observe differences in MG and non-escapes, this property can be used to distinguish the two.
184:
flexion is a hundredth of a second (10 milliseconds). Finally, the caridoid escape reflex requires that neurons be able to complete the arduous task of synchronizing the flexion of several abdominal segments. The speed, coordination, and decisiveness of the process seem to be the main attributes to its success.
382:
LGI itself is modulated by the behavior. Feeding increases the threshold for the tail flip, and even when the tail is tapped, no spikes are initiated. As a result, it takes an especially strong stimulus to elicit the tail flip. This prevents the crayfish from going hungry unless it is absolutely necessary.
408:
Repeated tapping of the abdomen leads to habituation of the tail flip mechanism. However, self–habituation is prevented by command neuron–derived inhibition because when a tail flip is begun, the mechanisms that induce habituation are repressed. The habituation occurs at the level of the A type and C
346:
of the MRO are located on the dorsal side of the abdomen, each spanning the joint of two segments. They are innervated by proprioreceptors that detect the stretch in extensor muscles when the flexors are contracted. They then fire phasically or tonically. It then excites the motor neurons in the fast
295:
Non-giant escape circuitry was found to be activated more during frontal attacks, but was rarely involved in the initial escape during a rear attack. Non-giant escape is often used after an initial giant interneuron-mediated tail flip. Compared to MG escape, the potentials produced in the MoG neurons
390:
When a crayfish is held by its carapace either in the water or in the air, it does not respond with a tail flip when it receives sensory stimuli that would normally elicit the response. This inhibitory behavior does not seem to be mediated by the abdomen. When the thorax-abdomen junction is severed,
332:
A number of processes are inhibited once the initial flexion has begun. Further LGI spiking, influx of sensory information from the tail, MoG spiking and FF contraction are all inhibited so that a flexion and extension cycle can be completed before another flexion begins. Long IPSPs are generated to
312:
The LGI is one of the few examples of a true command neuron. It passes the necessity and sufficiency principle put forward by
Kupfermann and Weiss. Wiersma's initial experiments showed that direct stimulation of the LGI was a sufficient release for the tail flip. In 1981, it was Olson and Krasne who
245:
known as the lateral giant neurons and the medial giant neurons. These interneurons play important roles in escape swimming. Their large diameter allows for rapid conduction since there is less current leakage. Their projections extend through the third root in each ganglion, and
Furshpan and Potter
421:
Changes in social status should correlate with changes in serotonin levels, resulting in changes in the escape strategies used by the crayfish. However this is only true when submissive crayfish become dominant, and not the reverse. The neuromodulatory processes of facilitation and inhibition seem
394:
The tail flip normally induces inhibition in an absolute fashion, such that it takes precedence over all the other tasks the segments are performing. However, during these situations, the effectiveness of the tail flip is reduced, so the inhibitory processes need to occur in a relative manner and a
291:
Non-giant escape often occurs during situations where lateral or medial giant-mediated escape may not be beneficial or during times where those behaviors are suppressed. This allows the crayfish to attack offenders, escape during feeding, or wriggle free when it has been restrained by the carapace.
200:
that reflects muscular and neural segmentation. The anterior portion of the crayfish is the cephalothorax region. The region rostral to the cephalic groove, which separates the head and thorax region, is characterized by the presence of eyes, antennae and claws while the region caudal contains four
381:
Crayfish often find themselves in a conflicting situation where they are performing the highly motivated behavior of feeding when they suddenly receive a tail flip stimulus. Often, the crayfish will not perform a tail flip in this situation. This is because when a crayfish is actively feeding, the
324:
The LGI receives its sensory input from the hair-like projections found on the edge of the tail fan. The sensory information is sent from bipolar receptors connected to the hair that are directionally sensitive. The path of the signal varies from here depending on the strength of the input. In the
287:
Less abrupt or gradually intensifying stimuli evoke locomotion behavior similar to that seen in giant interneuron-mediated behaviors. However, the movements are less stereotyped and do not appear to occur in an all or nothing manner. They are so named because they lack the involvement of the giant
183:
are used like a paddle to propel the crustacean away from harm using powerful abdominal flexions. The entire process occurs in a fraction of a second as movements are generated within two hundredths of a second (20 milliseconds) from the original trigger stimulus and the period of latency after a
341:
When the LGI-mediated inhibition subsides, the re-extension process begins. This process is mediated by the MRO and tail fan hair receptors, which were inhibited during the flexion portion of the escape behavior. The MRO is responsible for the initiation of extension and the inhibition of flexion
328:
The convergence many inputs on the LGI and divergence of few signals show how the LGI functions as a decision making coincidence detector. It takes the information from both the alpha and the beta pathways and if the timing of the spikes is synchronous, the behavior is produced. If the timing is
178:
depends on the region of the crayfish that is stimulated but all forms require abdominal contractions. When a strong, unpleasant tactile stimulus is presented, such as a burst of water or the prod of a probe, a stereotyped behavior occurs in which the muscular tail and wide tail fan region of the
1247:
provides an idea of the organization of the segmental ganglia in the tail of the crayfish. The second diagram on the page is a transverse section through the tail that highlights the positions of the LGI, MGI and non-giant neurons. At the bottom of the page it also has diagrams of the tail flips
443:
that diverged from the crayfish lineage very early on. It is likely that this mechanism was replaced by the tail flip since the wide surface area of the tail made this behavior more selectively advantageous. This most likely occurred when the ancestral flexor motor neurons in each segment formed
359:
Non-giant-mediated responses are initiated after the tail flip, creating cycles of flexion followed by extension. This non-giant system is activated parallel to the LGI circuit when the hair cells receive input. However, this behavior has a longer delay that allows the onset of swimming to occur
430:
It has been hypothesized that the tail flip is derived from an ancient limb protraction driven (as opposed to tail flexion-driven) mechanism. This is because the SGs appear to be modified limb motor neurons whose peripheral axons affect the legs and swimmerets, but end blindly without any known
417:
The feedback from social situations affect a crayfish's ability to perform a tail flip. Serotonin levels are affected by social status. High levels are associated with aggressive behavior and a reduction in the frequency of tail flips performed. This is because the excitability of the LGI is
250:
account for the speed of the escape mechanism and display some features of chemical synapses such as LTP and LTD. Variations in escape response characteristic depend on the location where the crayfish body is prodded or attacked and also depend on which of the giant neurons is stimulated.
278:
Unfortunately, this variant of the tail-flip escape has not been extensively studied. Further studies should focus on this escape variant, paying special attention to exactly how visual information is processed and then converted into neuronal signals that produce a tail flip response.
133:(MG or MGI) would result in the execution of a tail flip. Wiersma and K. Ikeda then proposed the term "command neuron" in their 1964 publication, and applied it to the giant interneuron's ability to command the expression of the escape response. This was the first description of a
329:
asynchronous, the later input is blocked by reducing the driving force of the signal and increasing the threshold voltage. The early synapse also causes current to leak through the active synapse, resulting in weak EPSPs that are ultimately unable to generate the tail flip.
232:
that innervate the extensor muscles, while the third root contains only motor neuron projections that extend into the flexor muscles. The last segment contains the fusion of two ganglia. The ganglia here also receive sensory input from the sensitive hairs on the tail fan.
84:
that make the crustacean quickly swim backwards through the water and away from danger. The type of response depends on the part of the crustacean stimulated, but this behavior is complex and is regulated both spatially and temporally through the interactions of several
313:
showed that the LGI fulfilled the condition of necessity, because when spikes were inhibited in the LGI through introducing a hyperpolarizing current, no motor activity was produced in response to a stimulus that would have originally elicited the tail flip.
399:
of the LGI. It is most likely that during restraint or feeding, the inhibitory process is mediated by inhibitory synaptic input on distal dendrites of the LGI . As a result, these signals would have to compete with other inputs for the control of the LGI.
434:
It has been speculated that the ancestral escape mechanism was most likely a backwards jump due to the simultaneous protraction of the legs driven by the ancestors of the Giant Fibers. This behavior was probably similar to the escape system found in a
372:
The production of a tail flip is not always beneficial. Evolution has allowed the crayfish to be more flexible by presenting several control systems that will prevent the tail flip in situations where it will most likely be unnecessary or ineffective.
448:
mechanism seems to be similar to this ancestral state because a large diameter axon in the dorsal nerve chord facilitates limb promoter motor neurons. This seems to match the ancestral condition, but it is not known for sure whether the circuitry is
223:
run down the entire length of the abdomen and communicate with one another through projections. The first five abdominal segments each have their own ganglion, that contains three roots with outward projections. The first has mixed sensory and
304:
The lateral giant (LG)-mediated escape mechanism is the most extensively analyzed form of the tail flip. The LG is not actually one neuron, but rather a group of closely associated neurons arranged end to end and connected by
124:
present in the tail were responsible for the reaction. The aforementioned neuronal fibres consist of a pair of lateral giant interneurons and a pair of medial giant interneurons, and
Wiersma found that stimulating just one
236:
Each ganglion contains the body of one motor giant neuron (MoG), powerful and large bodied motor neurons whose projections innervate the five fast flexor (FF) muscles found in a segment and interact with them through
1248:
caused by stimulation of the LGI (on the left half of the diagram) and the MGI (on the right half of the diagram). This accurate diagram appears to be similar to that found in Wine and Krasne's 1972 publication.
1209:
Roberts, A., Krasne, F. B., Hagiwara, G., Wine, J. J., and
Krarner, A. P. (1982) "Segmental giant: Evidence for a driver neuron interposed between command and motor neurons in the crayfish escape system".
409:
type interneurons, which experience synaptic depression. The habituation process is also mediated further up the circuit through the buildup of tonic inhibition, brought on by the repeated stimulation.
431:
function. It is known that another effect of Giant Fiber excitation is limb promotion which suggests that the premotor limb interneurons may be ancestors of the Giant Fibers.
170:
it was discovered that the tail-flip mechanism is characterized by a decisive, all-or-nothing quality that inhibits all unnecessary behaviors while generating a
641:
1103:
Vu, E. T.; Lee, S. C.; Krasne, F. B. (1993), "The mechanism of tonic inhibition of crayfish escape behavior: distal inhibition and its functional",
1150:
1290:
1221:
Wine, J.J. and
Hagiwara, G. (1977) "Crayfish Escape Behavior I. The Structure of Efferent and Afferent Neurons Involved in Abdominal Extension".
1241:
includes information on the biology and conservation of lobsters. The majority can be applied to crayfish due to common ancestry and homology.
785:
1318:
941:
863:
632:
535:
1335:
395:
tail flip should occur only if absolutely necessary. During absolute inhibition is directed by more proximal synapses controlling the
246:
found that the synapses they subsequently made with the MoG passed depolarizing currents in a direct and unidirectional manner. These
351:
detect the resulting movement caused by the tail flip when activated, they would fire and excite the fast extensor motor neurons.
705:
Wiersma, C. A.; Ikeda, K. (1964), "Interneurons
Commanding Swimmeret Movements in the Crayfish, Procambarus Clarki (girard)",
1534:
144:
This concept was further fleshed out with more specific and stringent conditions in 1972 when
Kupfermann and Weiss published
1251:
993:
Kramer, A. P.; Krasne, F. B. (1984), "Crayfish escape behavior: production of tailflips without giant fiber activity",
1283:
959:
Reichert, H.; Wine, J. J. (1983), "Coordination of lateral giant and non-giant systems in crayfish escape behavior",
638:
1539:
1524:
1340:
530:
510:
126:
347:
extensor muscles while directly exciting an inhibitory neuron that prevents contractions in the FF muscles. The
1602:
515:
130:
1629:
1397:
76:
The reaction, most extensively researched in crayfish, allows crustaceans to escape predators through rapid
396:
1624:
1590:
1554:
1313:
1276:
1370:
1147:
272:
1238:
1032:"Escape behavior and escape circuit activation in juvenile crayfish during prey-predator interactions"
111:
1447:
1380:
1360:
1355:
525:
171:
48:
1514:
1452:
1437:
1402:
824:
540:
162:
116:
103:
1477:
1407:
1375:
1350:
976:
816:
756:
450:
306:
247:
666:"Giant Nerve Fiber System of the Crayfish. A Contribution to Comparative Physiology of Synapse"
1498:
1263:
1199:
1130:
1053:
1010:
937:
911:
859:
808:
722:
685:
628:
1564:
1559:
1472:
1191:
1120:
1112:
1086:
1043:
1002:
968:
903:
800:
748:
714:
677:
361:
316:
The LGI contains the following three phases, occurring over the span of about 100 ms:
238:
44:
1529:
1432:
1427:
1154:
645:
175:
40:
19:
1031:
665:
97:
1549:
1422:
1417:
1365:
1125:
1116:
932:
Neuroethology : an introduction to the neurophysiological fundamentals of behavior
930:
520:
160:
have researched this behavior extensively for over fifty years in the crayfish species
138:
134:
1071:
804:
786:"Fifty years of a command neuron: the neurobiology of escape behavior in the crayfish"
1618:
1412:
1345:
1299:
1176:
760:
718:
436:
157:
980:
1569:
1519:
1457:
888:
820:
229:
264:
The medial giants innervate the entire ventral nerve chord. Their cell bodies and
1493:
1462:
1442:
225:
197:
121:
1244:
1467:
752:
54:
1006:
681:
625:
Behavioral neurobiology : the cellular organization of natural behavior
348:
77:
1057:
915:
812:
726:
689:
268:
begin in the brain and collect sensory information presented by visual and
1203:
1134:
1090:
1014:
141:
of a neuron could precipitate complex innate behaviors in some organisms.
1330:
1255:
1195:
505:
265:
213:
193:
70:
51:
565:
Krasne, F. B.; Wine, J. J. (1987), "Evasion responses of the crayfish",
972:
907:
488:
343:
220:
58:
1048:
1324:
478:
465:
216:
209:
202:
180:
86:
81:
66:
205:. The abdominal segments contain swimmerets, which aid in swimming.
739:
Kupfermann, I.; Weiss, K.R. (1978), "The command neuron concept",
269:
101:
The tail-flip escape behavior was first described in the crayfish
96:
62:
18:
1268:
242:
1272:
1030:
Herberholz, Jens; Sen, Marjorie M.; Edwards, Donald H. (2004),
1177:"Command Interneurons in the Crayfish Central Nervous System"
471:
208:
The anterior five segments of the crayfish house the massive
486:, which is the root of the name of the taxonomic infraorder
228:
innervating swimmerets while the second has sensory and
1072:"The Structural Basis of an Innate Behavioural Pattern"
93:
Discovery of the first command neuron-mediated behavior
889:"The Organization of Escape Behaviour in the Crayfish"
784:
Edwards, D. H.; Heitler, W. J.; Krasne, F. B. (1999),
444:
connections with one of these limb motor neurons. The
196:
crustaceans, the crayfish possesses a hard, segmented
114:
first described the tail-flip escape in the crayfish
627:, Sunderland, Mass.: Sinauer Associates Publishers,
1507:
1486:
1390:
1306:
929:
1025:
1023:
858:, Sunderland, Massachusetts: Sinauer Associates,
492:, a large group of crustaceans also known as the
954:
952:
1264:Cornell University's Neuroethology Course page
1284:
779:
777:
775:
773:
771:
769:
700:
698:
659:
657:
655:
653:
241:. The ganglia also contain two sets of giant
137:-mediated behavior and it indicated that the
8:
618:
616:
614:
612:
610:
608:
606:
604:
602:
600:
598:
596:
560:
558:
556:
849:
847:
845:
843:
594:
592:
590:
588:
586:
584:
582:
580:
578:
576:
426:Evolution of the tail flip escape mechanism
156:Behavioral neurobiologists in the field of
1291:
1277:
1269:
882:
880:
878:
876:
874:
1175:Atwood, H. L.; Wiersma, C. A. G. (1967),
1124:
1047:
368:Modulation and plasticity of the response
1223:The Journal of Comparative Physiology-A
552:
364:for physical and temporal maintenance.
639:Chapter 7: Escape Behavior in Crayfish
23:Animated representation of lobstering.
16:Innate escape mechanism by crustaceans
536:Coincidence detection in neurobiology
342:through a reflex like mechanism. The
255:Response variants and their circuitry
7:
1597:
961:Journal of Comparative Physiology A
887:Wine, J. J.; Krasne, F. B. (1972),
1117:10.1523/JNEUROSCI.13-10-04379.1993
14:
567:Aims and Methods in Neuroethology
174:for escape swimming. The type of
1596:
1585:
1584:
1341:Central pattern generator (CPG)
1184:Journal of Experimental Biology
1079:Journal of Experimental Biology
1036:Journal of Experimental Biology
896:Journal of Experimental Biology
80:flexions that produce powerful
1535:Frog hearing and communication
1:
805:10.1016/S0166-2236(98)01340-X
300:Lateral giant-mediated escape
719:10.1016/0010-406X(64)90153-7
466:
260:Medial giant-mediated escape
936:, Berlin: Springer-Verlag,
1648:
1540:Infrared sensing in snakes
1525:Jamming avoidance response
1212:Journal of Neurophysiology
995:Journal of Neurophysiology
928:Ewert, Jörg-Peter (1980),
854:Camhi, Jeffrey M. (1984),
670:Journal of Neurophysiology
664:Wiersma, C. A. G. (1947),
472:
464:is derived from the Greek
146:The Command Neuron Concept
1580:
753:10.1017/S0140525X00059057
623:Carew, Thomas J. (2000),
531:Central pattern generator
511:Lateral giant interneuron
476:(pronounced "kah-RISS"),
127:lateral giant interneuron
120:and noted that the giant
1545:Caridoid escape reaction
1254:presents information on
1239:The Lobster Conservatory
1007:10.1152/jn.1984.52.2.189
516:Medial giant interneuron
131:medial giant interneuron
29:caridoid escape reaction
1398:Theodore Holmes Bullock
1245:Neural and tail anatomy
1105:Journal of Neuroscience
793:Trends in Neurosciences
682:10.1152/jn.1947.10.1.23
1555:Surface wave detection
397:spike-initiating zones
320:1 – Flexion production
166:. Based on studies of
152:Research with crayfish
107:
24:
1371:Anti-Hebbian learning
1091:10.1242/jeb.112.1.283
644:June 9, 2011, at the
100:
22:
1448:Bernhard Hassenstein
1381:Ultrasound avoidance
1356:Fixed action pattern
1319:Coincidence detector
1196:10.1242/jeb.46.2.249
1161:online Greek lexicon
1070:Wine, J. J. (1984),
707:Comp Biochem Physiol
526:Fixed action pattern
172:fixed action pattern
1515:Animal echolocation
1453:Werner E. Reichardt
1403:Walter Heiligenberg
541:Pain in crustaceans
307:electrical synapses
248:electrical synapses
163:Procambarus clarkii
129:(LG or LGI) or one
117:Procambarus clarkii
104:Procambarus clarkii
1478:Fernando Nottebohm
1376:Sound localization
1351:Lateral inhibition
1153:2018-09-04 at the
973:10.1007/BF00610337
908:10.1242/jeb.56.1.1
386:Carapace restraint
108:
25:
1612:
1611:
1499:Slice preparation
1361:Krogh's Principle
1336:Feature detection
1252:Lobster Body Plan
1169:Suggested reading
1111:(10): 4379–4393,
1049:10.1242/jeb.00992
1042:(11): 1855–1863,
943:978-0-387-09790-9
865:978-0-87893-075-3
634:978-0-87893-084-5
239:chemical synapses
1637:
1600:
1599:
1588:
1587:
1565:Mechanoreception
1560:Electroreception
1473:Masakazu Konishi
1438:Jörg-Peter Ewert
1293:
1286:
1279:
1270:
1206:
1181:
1162:
1144:
1138:
1137:
1128:
1100:
1094:
1093:
1076:
1067:
1061:
1060:
1051:
1027:
1018:
1017:
990:
984:
983:
956:
947:
946:
935:
925:
919:
918:
893:
884:
869:
868:
851:
838:
837:
836:
835:
829:
823:, archived from
790:
781:
764:
763:
736:
730:
729:
702:
693:
692:
661:
648:
637:
620:
571:
570:
562:
475:
474:
469:
362:sensory feedback
337:2 – Re-extension
283:Non-giant escape
219:. Six abdominal
188:Anatomy involved
112:C. A. G. Wiersma
31:, also known as
1647:
1646:
1640:
1639:
1638:
1636:
1635:
1634:
1615:
1614:
1613:
1608:
1576:
1530:Vision in toads
1503:
1482:
1433:Erich von Holst
1428:Karl von Frisch
1386:
1302:
1297:
1235:
1179:
1174:
1171:
1166:
1165:
1155:Wayback Machine
1145:
1141:
1102:
1101:
1097:
1074:
1069:
1068:
1064:
1029:
1028:
1021:
992:
991:
987:
958:
957:
950:
944:
927:
926:
922:
891:
886:
885:
872:
866:
853:
852:
841:
833:
831:
827:
788:
783:
782:
767:
741:Behav Brain Sci
738:
737:
733:
704:
703:
696:
663:
662:
651:
646:Wayback Machine
635:
622:
621:
574:
564:
563:
554:
549:
502:
494:caridean shrimp
459:
428:
415:
406:
388:
379:
370:
357:
339:
322:
302:
285:
262:
257:
190:
176:escape response
154:
95:
41:escape behavior
39:, is an innate
17:
12:
11:
5:
1645:
1644:
1641:
1633:
1632:
1630:Marine biology
1627:
1617:
1616:
1610:
1609:
1607:
1606:
1594:
1581:
1578:
1577:
1575:
1574:
1573:
1572:
1562:
1557:
1552:
1550:Vocal learning
1547:
1542:
1537:
1532:
1527:
1522:
1517:
1511:
1509:
1505:
1504:
1502:
1501:
1496:
1490:
1488:
1484:
1483:
1481:
1480:
1475:
1470:
1465:
1460:
1455:
1450:
1445:
1440:
1435:
1430:
1425:
1423:Donald Kennedy
1420:
1418:Donald Griffin
1415:
1410:
1408:Niko Tinbergen
1405:
1400:
1394:
1392:
1388:
1387:
1385:
1384:
1378:
1373:
1368:
1366:Hebbian theory
1363:
1358:
1353:
1348:
1343:
1338:
1333:
1328:
1321:
1316:
1310:
1308:
1304:
1303:
1298:
1296:
1295:
1288:
1281:
1273:
1267:
1266:
1261:
1260:
1259:
1249:
1234:
1233:External links
1231:
1230:
1229:
1219:
1207:
1190:(2): 249–261,
1170:
1167:
1164:
1163:
1139:
1095:
1085:(1): 283–319,
1062:
1019:
1001:(2): 189–211,
985:
948:
942:
920:
870:
864:
839:
799:(4): 153–161,
765:
731:
694:
649:
633:
572:
551:
550:
548:
545:
544:
543:
538:
533:
528:
523:
521:Command neuron
518:
513:
508:
501:
498:
458:
455:
427:
424:
414:
411:
405:
402:
387:
384:
378:
375:
369:
366:
356:
353:
338:
335:
321:
318:
301:
298:
284:
281:
261:
258:
256:
253:
189:
186:
153:
150:
139:depolarization
135:command neuron
94:
91:
15:
13:
10:
9:
6:
4:
3:
2:
1643:
1642:
1631:
1628:
1626:
1625:Neuroethology
1623:
1622:
1620:
1605:
1604:
1595:
1593:
1592:
1583:
1582:
1579:
1571:
1568:
1567:
1566:
1563:
1561:
1558:
1556:
1553:
1551:
1548:
1546:
1543:
1541:
1538:
1536:
1533:
1531:
1528:
1526:
1523:
1521:
1518:
1516:
1513:
1512:
1510:
1506:
1500:
1497:
1495:
1492:
1491:
1489:
1485:
1479:
1476:
1474:
1471:
1469:
1466:
1464:
1461:
1459:
1456:
1454:
1451:
1449:
1446:
1444:
1441:
1439:
1436:
1434:
1431:
1429:
1426:
1424:
1421:
1419:
1416:
1414:
1413:Konrad Lorenz
1411:
1409:
1406:
1404:
1401:
1399:
1396:
1395:
1393:
1389:
1382:
1379:
1377:
1374:
1372:
1369:
1367:
1364:
1362:
1359:
1357:
1354:
1352:
1349:
1347:
1346:NMDA receptor
1344:
1342:
1339:
1337:
1334:
1332:
1329:
1327:
1326:
1322:
1320:
1317:
1315:
1312:
1311:
1309:
1305:
1301:
1300:Neuroethology
1294:
1289:
1287:
1282:
1280:
1275:
1274:
1271:
1265:
1262:
1257:
1253:
1250:
1246:
1243:
1242:
1240:
1237:
1236:
1232:
1227:
1224:
1220:
1217:
1213:
1208:
1205:
1201:
1197:
1193:
1189:
1185:
1178:
1173:
1172:
1168:
1160:
1157:
1156:
1152:
1149:
1143:
1140:
1136:
1132:
1127:
1122:
1118:
1114:
1110:
1106:
1099:
1096:
1092:
1088:
1084:
1080:
1073:
1066:
1063:
1059:
1055:
1050:
1045:
1041:
1037:
1033:
1026:
1024:
1020:
1016:
1012:
1008:
1004:
1000:
996:
989:
986:
982:
978:
974:
970:
966:
962:
955:
953:
949:
945:
939:
934:
933:
924:
921:
917:
913:
909:
905:
901:
897:
890:
883:
881:
879:
877:
875:
871:
867:
861:
857:
856:Neuroethology
850:
848:
846:
844:
840:
830:on 2007-06-11
826:
822:
818:
814:
810:
806:
802:
798:
794:
787:
780:
778:
776:
774:
772:
770:
766:
762:
758:
754:
750:
746:
742:
735:
732:
728:
724:
720:
716:
713:(4): 509–25,
712:
708:
701:
699:
695:
691:
687:
683:
679:
675:
671:
667:
660:
658:
656:
654:
650:
647:
643:
640:
636:
630:
626:
619:
617:
615:
613:
611:
609:
607:
605:
603:
601:
599:
597:
595:
593:
591:
589:
587:
585:
583:
581:
579:
577:
573:
568:
561:
559:
557:
553:
546:
542:
539:
537:
534:
532:
529:
527:
524:
522:
519:
517:
514:
512:
509:
507:
504:
503:
499:
497:
495:
491:
490:
485:
481:
480:
468:
463:
456:
454:
452:
447:
442:
438:
437:mantis shrimp
432:
425:
423:
419:
413:Social status
412:
410:
403:
401:
398:
392:
385:
383:
376:
374:
367:
365:
363:
354:
352:
350:
345:
344:muscle fibers
336:
334:
330:
326:
319:
317:
314:
310:
308:
299:
297:
293:
289:
282:
280:
276:
274:
271:
267:
259:
254:
252:
249:
244:
240:
234:
231:
230:motor neurons
227:
222:
218:
215:
211:
206:
204:
199:
195:
187:
185:
182:
177:
173:
169:
165:
164:
159:
158:neuroethology
151:
149:
147:
142:
140:
136:
132:
128:
123:
119:
118:
113:
106:
105:
99:
92:
90:
88:
83:
79:
74:
72:
68:
64:
60:
56:
53:
50:
46:
42:
38:
37:tail-flipping
34:
30:
21:
1601:
1589:
1570:Lateral line
1544:
1520:Waggle dance
1458:Eric Knudsen
1323:
1225:
1222:
1215:
1211:
1187:
1183:
1158:
1146:
1142:
1108:
1104:
1098:
1082:
1078:
1065:
1039:
1035:
998:
994:
988:
964:
960:
931:
923:
899:
895:
855:
832:, retrieved
825:the original
796:
792:
744:
740:
734:
710:
706:
676:(1): 23–38,
673:
669:
624:
566:
493:
487:
483:
477:
461:
460:
445:
440:
433:
429:
420:
416:
407:
393:
389:
380:
371:
358:
355:3 – Swimming
340:
331:
327:
323:
315:
311:
303:
294:
290:
286:
277:
263:
235:
226:motor nerves
207:
191:
167:
161:
155:
145:
143:
122:interneurons
115:
109:
102:
75:
36:
32:
28:
26:
1494:Patch clamp
1463:Eric Kandel
1443:Franz Huber
1314:Feedforward
967:(1): 3–15,
902:(1): 1–18,
198:exoskeleton
192:Like other
55:crustaceans
1619:Categories
1468:Nobuo Suga
1383:in insects
1228:: 145–172.
1218:: 761–781.
834:2008-11-10
547:References
451:homologous
349:hair cells
168:P. clarkii
49:freshwater
33:lobstering
761:143708768
457:Etymology
266:dendrites
110:In 1946,
78:abdominal
1591:Category
1331:Instinct
1307:Concepts
1258:anatomy.
1151:Archived
1058:15107440
981:41122772
916:21046844
813:10203852
747:: 3–39,
727:14206963
690:20279137
642:Archived
506:Crayfish
500:See also
462:Caridoid
404:Learning
214:extensor
71:crayfish
59:lobsters
57:such as
1603:Commons
1508:Systems
1487:Methods
1256:decapod
1204:6033993
1159:ΛΟΓΕΙΟΝ
1135:8410194
1126:6576376
1015:6090603
821:1197778
569:: 10–45
489:Caridea
446:Squilla
441:Squilla
439:called
273:stimuli
270:tactile
221:ganglia
217:muscles
194:decapod
87:neurons
82:thrusts
52:eucarid
1391:People
1325:Umwelt
1202:
1133:
1123:
1056:
1013:
979:
940:
914:
862:
819:
811:
759:
725:
688:
631:
479:shrimp
377:Eating
210:flexor
203:telson
181:telson
67:shrimp
45:marine
1180:(PDF)
1148:καρίς
1075:(PDF)
977:S2CID
892:(PDF)
828:(PDF)
817:S2CID
789:(PDF)
757:S2CID
484:prawn
473:καρίς
467:karis
243:axons
63:krill
1200:PMID
1131:PMID
1054:PMID
1011:PMID
938:ISBN
912:PMID
860:ISBN
809:PMID
723:PMID
686:PMID
629:ISBN
212:and
69:and
47:and
27:The
1226:121
1192:doi
1121:PMC
1113:doi
1087:doi
1083:112
1044:doi
1040:207
1003:doi
969:doi
965:153
904:doi
801:doi
749:doi
715:doi
678:doi
482:or
470:or
43:in
35:or
1621::
1216:47
1214:.
1198:,
1188:46
1186:,
1182:,
1129:,
1119:,
1109:13
1107:,
1081:,
1077:,
1052:,
1038:,
1034:,
1022:^
1009:,
999:52
997:,
975:,
963:,
951:^
910:,
900:56
898:,
894:,
873:^
842:^
815:,
807:,
797:22
795:,
791:,
768:^
755:,
743:,
721:,
711:12
709:,
697:^
684:,
674:10
672:,
668:,
652:^
575:^
555:^
496:.
453:.
89:.
73:.
65:,
61:,
1292:e
1285:t
1278:v
1194::
1115::
1089::
1046::
1005::
971::
906::
803::
751::
745:1
717::
680::
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
