27:
251:, but innervates only about a dozen different pyramidal cells. In contrast, a single CA3 pyramidal cell receives input from about 50 different granule cells. It has been shown in rodents that the size of the mossy fiber projections can show large interindividual variations, which are to a large part heritable. In addition, these variations show strong correlations with different types of behavior, mainly, but not exclusively,
31:
Stippled area: strata oriens (OR) and radiatum (RD). Hatched area: stratum lacunosum-moleculare (LM). CA1, subregion of the hippocampus without mossy fibers; FI, fimbria hippocampi; FD, fascia dentata; OL and ML, outer and middle molecular layers of the fascia dentata; SG, supragranular layer; GC, granular cells.
238:
have been shown to be mediated by sodium channels that receive input from neuroglia. The entorhinal cortex also projects directly to CA3, suggesting that the mossy fiber pathway may be functionally similar to the perforant pathway although microcircuits within the dentate gyrus give the mossy fiber
194:
with 25–35 synaptic connections and filopodial extensions with 12–17 make up a significant portion of total granule cell synaptic terminals and are mainly responsible for the excitation of GABAergic interneurons. The type of synaptic terminal expressed therefore dictates the downstream targeting of
30:
Diagram of a Timm-stained cross-section of the mouse hippocampus. The hippocampal subregion CA3–CA4 is indicated in black, stippled, and hatched areas. Black areas: suprapyramidal (SP), intra- and infrapyramidal (IIP) and hilar (CA4) mossy fiber terminal fields originating from the dentate gyrus.
116:
neurotransmitters within mossy fiber terminals. GABAergic and glutamatergic co-localization in mossy fiber boutons has been observed primarily in the developing hippocampus, but in adulthood, evidence suggests that mossy fiber synapses may alternate which neurotransmitter is released through
239:
pathway a more modulatory role. Projections to the dentate hilus are excitatory by nature and oppose the inhibitory effects of interneurons on hilar mossy cells. The result is an excitatory feedforward loop on mossy cells as a result of activation by the entorhinal cortex.
222:
that allow them to be modulated by both excitatory and inhibitory input from nearby glial cells. Axons from the entorhinal cortex synapse primarily on the dendritic spines of outer layer dentate granule cells. The entorhinal cortex passes sensory information from
186:. Hilar mossy cell activation is thought to be necessary for the proper function of these inhibitory basket cells on CA3 pyramidal cells, although evidence has shown that sodium channel receptors can regulate basket cell function as well.
189:
The three synaptic terminals types – mossy terminals, filopodial extensions, and en passant synaptic varicosities – differ in synaptic output. Large mossy terminals synapse with 11–15 different CA3 pyramidal cells and 7–12 mossy cells.
162:. Thorny excrescences also cover the proximal dendrites of mossy cells in the hilus. Hilar thorny excrescences are more dense and complex than those in CA3. It has been shown that the axons of granule cells from the dentate gyrus
137:
of hilar mossy cells and pyramidal cells in CA3. They form three morphologically different synaptic terminals, which include large mossy terminals, filopodial extensions within the mossy terminals, and small
195:
granule cells. The high convergence onto pyramidal cells and divergent projections onto interneurons suggests a primarily modulatory role for the mossy fiber pathway in the hippocampus.
234:
The mossy fiber pathway itself projects to CA3. Repetitive stimulation of its neurons leads to progressive use-dependent synaptic depression. These short-term changes in
933:
Danscher G, Zimmer J (February 1978). "An improved Timm sulphide silver method for light and electron microscopic localization of heavy metals in biological tissues".
890:
Gonzales RB, DeLeon Galvan CJ, Rangel YM, Claiborne BJ (February 2001). "Distribution of thorny excrescences on CA3 pyramidal neurons in the rat hippocampus".
636:
Sandler R, Smith AD (January 1991). "Coexistence of GABA and glutamate in mossy fiber terminals of the primate hippocampus: an ultrastructural study".
73:
The axons that make up the pathway emerge from the basal portions of the granule cells and pass through the hilus (or polymorphic cell layer) of the
26:
438:
Amaral DG, Scharfman HE, Lavenex P (2007). "The dentate gyrus: fundamental neuroanatomical organization (dentate gyrus for dummies)".
285:"Prenatal exposure to alcohol does not affect radial maze learning and hippocampal mossy fiber sizes in three inbred strains of mouse"
206:
The dentate gyrus receives excitatory projections from neurons in layer II of the entorhinal cortex as well as input from surrounding
1151:
455:
405:
903:
1087:"An analysis of entorhinal cortex projections to the dentate gyrus, hippocampus, and subiculum of the neonatal macaque monkey"
155:
78:
1320:
1035:"Short-Term Depression of Axonal Spikes at the Mouse Hippocampal Mossy Fibers and Sodium Channel-Dependent Modulation"
1266:"Learning spatial orientation tasks in the radial-maze and structural variation in the hippocampus in inbred mice"
1221:, Genthner-Grimm G, Schwegler H (2007). "A quantitative-genetic analysis of hippocampal variation in the mouse".
67:
264:
20:
1175:
Henze DA, Urban NN, Barrionuevo G (2000). "The multifarious hippocampal mossy fiber pathway: a review".
228:
1134:
Witter MP (2007). "The perforant path: projections from the entorhinal cortex to the dentate gyrus".
681:"Plasticity of the GABAergic phenotype of the "glutamatergic" granule cells of the rat dentate gyrus"
130:
86:
59:
1246:
1200:
958:
915:
661:
388:
Henze DA, Buzsáki G (2007). "Hilar mossy cells: functional identification and activity in vivo".
191:
159:
139:
1297:
1238:
1192:
1157:
1147:
1116:
1064:
1012:
950:
907:
872:
814:
763:
712:
653:
618:
564:
515:
471:
451:
411:
401:
370:
316:
179:
63:
1287:
1277:
1230:
1184:
1139:
1106:
1098:
1054:
1046:
1002:
992:
942:
899:
862:
852:
804:
794:
753:
743:
702:
692:
645:
608:
598:
554:
546:
505:
461:
443:
393:
360:
352:
306:
296:
252:
247:
In the mouse, a single mossy fiber projection may make as many as 37 contacts with a single
215:
90:
841:"GABAergic cells are the major postsynaptic targets of mossy fibers in the rat hippocampus"
679:
Gutiérrez R, Romo-Parra H, Maqueda J, Vivar C, Ramìrez M, Morales MA, Lamas M (July 2003).
1136:
The
Dentate Gyrus: A Comprehensive Guide to Structure, Function, and Clinical Implications
440:
The
Dentate Gyrus: A Comprehensive Guide to Structure, Function, and Clinical Implications
390:
The
Dentate Gyrus: A Comprehensive Guide to Structure, Function, and Clinical Implications
235:
198:
The synapses of the mossy fibers contain zinc, which can be stained with a Timm staining.
340:
1292:
1265:
1111:
1086:
1059:
1034:
1007:
980:
867:
857:
840:
809:
782:
758:
731:
707:
697:
680:
613:
586:
559:
534:
490:
466:
365:
311:
284:
248:
183:
175:
151:
1188:
1143:
447:
397:
70:
as they displayed varicosities along their lengths that gave them a mossy appearance.
1314:
219:
211:
126:
113:
82:
74:
55:
1204:
665:
341:"Local and Long-Range Circuit Connections to Hilar Mossy Cells in the Dentate Gyrus"
1250:
962:
799:
748:
51:
919:
535:"Distinct roles for dorsal CA3 and CA1 in memory for sequential nonspatial events"
1050:
981:"Structure, function, and plasticity of hippocampal dentate gyrus microcircuits"
356:
231:. The pathway allows sensory information to reach the hippocampus for encoding.
171:
167:
36:
150:
Mossy fibers form multiple synapses with the elaborate dendritic spines of CA3
1234:
1218:
210:. The unmyelinated granule cell axons of the mossy fiber pathway express both
105:
44:
997:
603:
783:"Is GABA co-released with glutamate from hippocampal mossy fiber terminals?"
224:
207:
133:. The pathway consists of varicose granule cell axons that terminate on the
109:
101:
1301:
1242:
1196:
1161:
1120:
1068:
1016:
911:
818:
767:
716:
622:
568:
519:
475:
415:
374:
320:
1282:
876:
657:
649:
301:
732:"GABAergic signaling at mossy fiber synapses in neonatal rat hippocampus"
134:
954:
904:
10.1002/1096-9861(20010212)430:3<357::aid-cne1036>3.0.co;2-k
946:
550:
163:
97:
1102:
510:
94:
587:"Hilar mossy cells of the dentate gyrus: a historical perspective"
25:
19:
For the major input axonal pathway bundle to the cerebellum, see
730:
Safiulina VF, Fattorini G, Conti F, Cherubini E (January 2006).
47:
392:. Progress in Brain Research. Vol. 163. pp. 199–216.
491:"Behavioral functions of the CA3 subregion of the hippocampus"
1138:. Progress in Brain Research. Vol. 163. pp. 43–61.
839:
Acsády L, Kamondi A, Sík A, Freund T, Buzsáki G (May 1998).
442:. Progress in Brain Research. Vol. 163. pp. 3–22.
283:
Sluyter F, Jamot L, Bertholet JY, Crusio WE (April 2005).
142:. Each of these synapse types is functionally distinct.
66:. These axons were first described as mossy fibers by
58:
that terminate on modulatory hilar mossy cells and in
108:. There is also evidence for co-localization of both
533:Farovik A, Dupont LM, Eichenbaum H (January 2010).
125:Mossy fibers in the hippocampus project from the
580:
578:
339:Sun Y, Grieco SF, Holmes TC, Xu X (2017-03-01).
834:
832:
830:
828:
89:data has indicated that some synapses contain
433:
431:
429:
427:
425:
8:
178:in the CA3 region, providing input from the
1085:Amaral DG, Kondo H, Lavenex P (May 2014).
334:
332:
330:
1291:
1281:
1110:
1058:
1006:
996:
866:
856:
808:
798:
757:
747:
706:
696:
612:
602:
558:
509:
465:
364:
310:
300:
81:of CA3. Granule cell synapses tend to be
275:
1264:Crusio WE, Schwegler H (April 2005).
1080:
1078:
1028:
1026:
974:
972:
166:with hilar mossy cells and GABAergic
62:(CA3), a region involved in encoding
7:
1091:The Journal of Comparative Neurology
892:The Journal of Comparative Neurology
638:The Journal of Comparative Neurology
158:. These complex spines are known as
1033:Ohura S, Kamiya H (January 2018).
858:10.1523/jneurosci.18-09-03386.1998
698:10.1523/jneurosci.23-13-05594.2003
156:stratum lucidum of the hippocampus
14:
585:Scharfman HE, Myers CE (2012).
117:activity-dependent regulation.
1270:Behavioral and Brain Functions
800:10.1523/JNEUROSCI.5019-12.2013
749:10.1523/JNEUROSCI.4493-05.2006
289:Behavioral and Brain Functions
1:
1189:10.1016/s0306-4522(00)00146-9
1144:10.1016/S0079-6123(07)63003-9
448:10.1016/S0079-6123(07)63001-5
398:10.1016/s0079-6123(07)63012-x
985:Frontiers in Neural Circuits
591:Frontiers in Neural Circuits
1051:10.1523/ENEURO.0415-17.2018
845:The Journal of Neuroscience
787:The Journal of Neuroscience
736:The Journal of Neuroscience
685:The Journal of Neuroscience
489:Kesner RP (November 2007).
357:10.1523/ENEURO.0097-17.2017
243:Role in learning and memory
1337:
1045:(1): ENEURO.0415–17.2018.
979:Jonas P, Lisman J (2014).
781:Caiati MD (January 2013).
351:(2): ENEURO.0097–17.2017.
85:(i.e. excitatory), though
18:
16:Pathway in the hippocampus
1235:10.1080/01677060701715827
1223:Journal of Neurogenetics
998:10.3389/fncir.2014.00107
604:10.3389/fncir.2012.00106
265:Mossy fiber (cerebellum)
21:mossy fiber (cerebellum)
225:neocortical structures
112:(i.e. inhibitory) and
68:Santiago Ramón y Cajal
32:
1283:10.1186/1744-9081-1-3
650:10.1002/cne.903030202
539:Learning & Memory
498:Learning & Memory
302:10.1186/1744-9081-1-5
229:hippocampal formation
29:
77:before entering the
60:Cornu Ammonis area 3
43:pathway consists of
1321:Hippocampus (brain)
216:glutamate receptors
160:thorny excrescences
93:elements including
947:10.1007/bf00496691
551:10.1101/lm.1616209
192:En passant boutons
146:Synaptic terminals
140:en passant boutons
87:immunohistological
33:
1103:10.1002/cne.23469
511:10.1101/lm.688207
184:perforant pathway
180:entorhinal cortex
64:short-term memory
1328:
1306:
1305:
1295:
1285:
1261:
1255:
1254:
1215:
1209:
1208:
1172:
1166:
1165:
1131:
1125:
1124:
1114:
1082:
1073:
1072:
1062:
1030:
1021:
1020:
1010:
1000:
976:
967:
966:
930:
924:
923:
887:
881:
880:
870:
860:
836:
823:
822:
812:
802:
778:
772:
771:
761:
751:
727:
721:
720:
710:
700:
676:
670:
669:
633:
627:
626:
616:
606:
582:
573:
572:
562:
530:
524:
523:
513:
495:
486:
480:
479:
469:
435:
420:
419:
385:
379:
378:
368:
336:
325:
324:
314:
304:
280:
253:spatial learning
174:before reaching
91:neuropeptidergic
50:projecting from
1336:
1335:
1331:
1330:
1329:
1327:
1326:
1325:
1311:
1310:
1309:
1263:
1262:
1258:
1217:
1216:
1212:
1174:
1173:
1169:
1154:
1133:
1132:
1128:
1097:(7): 1485–505.
1084:
1083:
1076:
1032:
1031:
1024:
978:
977:
970:
932:
931:
927:
889:
888:
884:
851:(9): 3386–403.
838:
837:
826:
780:
779:
775:
729:
728:
724:
678:
677:
673:
635:
634:
630:
584:
583:
576:
532:
531:
527:
493:
488:
487:
483:
458:
437:
436:
423:
408:
387:
386:
382:
338:
337:
328:
282:
281:
277:
273:
261:
245:
204:
176:pyramidal cells
152:pyramidal cells
148:
123:
79:stratum lucidum
24:
17:
12:
11:
5:
1334:
1332:
1324:
1323:
1313:
1312:
1308:
1307:
1256:
1229:(4): 197–208.
1210:
1167:
1152:
1126:
1074:
1022:
968:
935:Histochemistry
925:
882:
824:
773:
742:(2): 597–608.
722:
691:(13): 5594–8.
671:
628:
574:
525:
504:(11): 771–81.
481:
456:
421:
406:
380:
326:
274:
272:
269:
268:
267:
260:
257:
249:pyramidal cell
244:
241:
212:GABA receptors
203:
200:
147:
144:
122:
119:
15:
13:
10:
9:
6:
4:
3:
2:
1333:
1322:
1319:
1318:
1316:
1303:
1299:
1294:
1289:
1284:
1279:
1275:
1271:
1267:
1260:
1257:
1252:
1248:
1244:
1240:
1236:
1232:
1228:
1224:
1220:
1214:
1211:
1206:
1202:
1198:
1194:
1190:
1186:
1183:(3): 407–27.
1182:
1178:
1171:
1168:
1163:
1159:
1155:
1153:9780444530158
1149:
1145:
1141:
1137:
1130:
1127:
1122:
1118:
1113:
1108:
1104:
1100:
1096:
1092:
1088:
1081:
1079:
1075:
1070:
1066:
1061:
1056:
1052:
1048:
1044:
1040:
1036:
1029:
1027:
1023:
1018:
1014:
1009:
1004:
999:
994:
990:
986:
982:
975:
973:
969:
964:
960:
956:
952:
948:
944:
940:
936:
929:
926:
921:
917:
913:
909:
905:
901:
898:(3): 357–68.
897:
893:
886:
883:
878:
874:
869:
864:
859:
854:
850:
846:
842:
835:
833:
831:
829:
825:
820:
816:
811:
806:
801:
796:
793:(5): 1755–6.
792:
788:
784:
777:
774:
769:
765:
760:
755:
750:
745:
741:
737:
733:
726:
723:
718:
714:
709:
704:
699:
694:
690:
686:
682:
675:
672:
667:
663:
659:
655:
651:
647:
644:(2): 177–92.
643:
639:
632:
629:
624:
620:
615:
610:
605:
600:
596:
592:
588:
581:
579:
575:
570:
566:
561:
556:
552:
548:
544:
540:
536:
529:
526:
521:
517:
512:
507:
503:
499:
492:
485:
482:
477:
473:
468:
463:
459:
457:9780444530158
453:
449:
445:
441:
434:
432:
430:
428:
426:
422:
417:
413:
409:
407:9780444530158
403:
399:
395:
391:
384:
381:
376:
372:
367:
362:
358:
354:
350:
346:
342:
335:
333:
331:
327:
322:
318:
313:
308:
303:
298:
294:
290:
286:
279:
276:
270:
266:
263:
262:
258:
256:
254:
250:
242:
240:
237:
232:
230:
226:
221:
217:
213:
209:
201:
199:
196:
193:
187:
185:
181:
177:
173:
169:
165:
161:
157:
153:
145:
143:
141:
136:
132:
128:
127:dentate gyrus
120:
118:
115:
114:glutamatergic
111:
107:
103:
99:
96:
92:
88:
84:
83:glutamatergic
80:
76:
75:dentate gyrus
71:
69:
65:
61:
57:
56:dentate gyrus
53:
52:granule cells
49:
46:
42:
38:
28:
22:
1273:
1269:
1259:
1226:
1222:
1213:
1180:
1177:Neuroscience
1176:
1170:
1135:
1129:
1094:
1090:
1042:
1038:
988:
984:
941:(1): 27–40.
938:
934:
928:
895:
891:
885:
848:
844:
790:
786:
776:
739:
735:
725:
688:
684:
674:
641:
637:
631:
594:
590:
545:(1): 12–17.
542:
538:
528:
501:
497:
484:
439:
389:
383:
348:
344:
292:
288:
278:
246:
233:
218:along their
205:
197:
188:
182:through the
172:basket cells
168:interneurons
149:
124:
72:
45:unmyelinated
40:
34:
202:Projections
41:mossy fiber
37:hippocampus
271:References
236:plasticity
170:including
106:enkephalin
1219:Crusio WE
220:membranes
208:neuroglia
135:dendrites
110:GABAergic
102:dynorphin
1315:Category
1302:15916698
1276:(1): 3.
1243:18161583
1205:11663807
1197:10869836
1162:17765711
1121:24122645
1069:29468192
1017:25309334
912:11169473
819:23365214
768:16407558
717:12843261
666:42502573
623:23420672
569:20028733
520:18007020
476:17765709
416:17765720
375:28451637
321:15916699
295:(1): 5.
259:See also
100:such as
98:peptides
1293:1143776
1251:7334116
1112:4384686
1060:5820996
1008:4159971
991:: 107.
963:2771576
877:9547246
868:6792657
810:6619118
759:6674413
708:6741238
658:1672874
614:3572871
597:: 106.
560:2807176
467:2492885
366:5396130
312:1143778
227:to the
164:synapse
154:in the
121:Anatomy
54:in the
35:In the
1300:
1290:
1249:
1241:
1203:
1195:
1160:
1150:
1119:
1109:
1067:
1057:
1039:eNeuro
1015:
1005:
961:
953:
920:916356
918:
910:
875:
865:
817:
807:
766:
756:
715:
705:
664:
656:
621:
611:
567:
557:
518:
474:
464:
454:
414:
404:
373:
363:
345:eNeuro
319:
309:
95:opiate
39:, the
1247:S2CID
1201:S2CID
959:S2CID
955:76622
916:S2CID
662:S2CID
494:(PDF)
48:axons
1298:PMID
1239:PMID
1193:PMID
1158:PMID
1148:ISBN
1117:PMID
1065:PMID
1013:PMID
951:PMID
908:PMID
873:PMID
815:PMID
764:PMID
713:PMID
654:PMID
619:PMID
565:PMID
516:PMID
472:PMID
452:ISBN
412:PMID
402:ISBN
371:PMID
317:PMID
214:and
104:and
1288:PMC
1278:doi
1231:doi
1185:doi
1140:doi
1107:PMC
1099:doi
1095:522
1055:PMC
1047:doi
1003:PMC
993:doi
943:doi
900:doi
896:430
863:PMC
853:doi
805:PMC
795:doi
754:PMC
744:doi
703:PMC
693:doi
646:doi
642:303
609:PMC
599:doi
555:PMC
547:doi
506:doi
462:PMC
444:doi
394:doi
361:PMC
353:doi
307:PMC
297:doi
131:CA3
129:to
1317::
1296:.
1286:.
1272:.
1268:.
1245:.
1237:.
1227:21
1225:.
1199:.
1191:.
1181:98
1179:.
1156:.
1146:.
1115:.
1105:.
1093:.
1089:.
1077:^
1063:.
1053:.
1041:.
1037:.
1025:^
1011:.
1001:.
987:.
983:.
971:^
957:.
949:.
939:55
937:.
914:.
906:.
894:.
871:.
861:.
849:18
847:.
843:.
827:^
813:.
803:.
791:33
789:.
785:.
762:.
752:.
740:26
738:.
734:.
711:.
701:.
689:23
687:.
683:.
660:.
652:.
640:.
617:.
607:.
593:.
589:.
577:^
563:.
553:.
543:17
541:.
537:.
514:.
502:14
500:.
496:.
470:.
460:.
450:.
424:^
410:.
400:.
369:.
359:.
347:.
343:.
329:^
315:.
305:.
291:.
287:.
255:.
1304:.
1280::
1274:1
1253:.
1233::
1207:.
1187::
1164:.
1142::
1123:.
1101::
1071:.
1049::
1043:5
1019:.
995::
989:8
965:.
945::
922:.
902::
879:.
855::
821:.
797::
770:.
746::
719:.
695::
668:.
648::
625:.
601::
595:6
571:.
549::
522:.
508::
478:.
446::
418:.
396::
377:.
355::
349:4
323:.
299::
293:1
23:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
