373:
lining that has been suggested to be a microtubule organizing centre for the axostyle. The size of parabasal fibers decreases as they extend further past the apex, to the point where they cannot be observed in the mid-region or base of the cell. Parabasal fibers are densely concentrated in the cell's apex, and axostyles closely associated with the parabasal fibers also accumulate in this location. The fibrous ribbon is a long sheet that looks like an accordion, and connects all the basal bodies in an individual flagellar band. An individual fibrous ribbon is as long as the length of an individual flagellar body. KI fibers are named for their distinctive shape, and specifically link basal bodies in triplets. KI fibers can change shape, which also changes the distance between basal bodies and regulates how close or far they are from each other. The fibrous ribbon and KI fiber are thought to have a role in controlling cell shape by moving the flagellar bands. They also play roles in regulating the direction a
Holomastigotoides cell moves in, coordinating the beating of flagella, and assisting in accommodating large pieces of wood during phagocytosis.
37:
382:
the cortical cytoplasm extend along the entire length of the flagellar bands. Some axostyles follow the spiral arrangement of the flagellar bands and regulate the positions of the Golgi bodies and endoplasmic reticulum in the cell. Notably, flagellar bands 4 and 5 are specialized, and possess extensions into the cytoplasm that contain the poles of the cell's extra-nuclear mitotic spindle.
362:
are more difficult to distinguish. As basal bodies become more widely spaced further away from the cell apex, the fiber types are also easier to distinguish. Basal bodies transition into flagella distally, and the transition point is indicated by a transition plate. An axosome is found between the transition plate and the central microtubules of an individual flagellum.
509:, telophase has been observed in greater detail. Telophase occurs via the separation and coiling of flagellar band. While this flagellar band coils, it pulls a daughter nucleus to the basal end of the cell. The number of flagellar bands in a daughter cell is determined by duplication of basal bodies at the end of cell division.
504:
has been observed to reproduce through asexual division. During cell division, the nucleus and chromosomes elongate longitudinally. A constriction forms in the middle of the nucleus until two daughter nuclei are produced, effectively splitting the chromosomes in half so that each daughter nucleus has
497:
can easily be distinguished due to size, as one will be shorter than the other. As the chromosomes replicate, they uncoil and appear to extend in length. After replication, the sister chromatids re-coil and shorten before separating and pairing with their homologues. Chromosomes have been observed to
381:
Axostyles can be located along the entire length of a flagellar band. They can extend from the cytoplasm to the cell base and surround the nucleus. They can also be found in the cortical cytoplasm, which is the cytoplasm that falls between the plasma membrane and flagellar basal bodies. Axostyles in
430:
cells with energy. The hydrogenosomes are located either between the plasma membrane and flagellar basal bodies or dispersed throughout the cytoplasm. They are thought to accumulate near the basal bodies to support high energy demands of the flagella, and have been observed to divide independently.
368:
also possesses parabasal bodies, as is characteristic of parabasalids. The parabasal bodies consist of a Golgi body and a parabasal fiber, and are closely associated with the basal bodies of the flagella. Golgi bodies have been observed to overlap with parabasal fibers near the base of the nucleus.
344:
its characteristic, highly flagellated appearance. The basal bodies of a flagellar band are linked by a fiber system that consists of three different fiber types. Each flagellar band is associated with an axostyle, endoplasmic reticulum, and Golgi bodies. The high density of external flagella helps
339:
is attributed to the arrangement of its flagellar bands in a spiral formation around the cell. The flagellar bands originate from the anterior apex of the cell and spiral posteriorly in progressively larger spirals, wrapping around the circumference of the cell. An individual flagellar band is made
461:
is unique in that it remains in the cell during most of the cell cycle, along with the flagella. Spindle poles are present to maintain spindle microtubules while the mitotic spindle is present. This is possible because cytoplasmic microtubules and mitotic microtubules have different origins in the
361:
Near the anterior apex of the cell, the basal bodies are arranged tightly together within the flagellar bands, to such an extent that some basal bodies will overlap with each other. The fiber system associated with the basal bodies is also compressed in this apical region, and thus the fiber types
401:
cells, there is a high concentration of centrin at the apex of the cell associated with the parabasal fibers, the flagellar bands, and the mitotic spindle. As these are sites where changes in cell shape and movement are initiated, this implies a possible role of centrin in controlling cell shape,
372:
The basal bodies of a flagellar band are linked by a fiber system, which consists of the parabasal fiber, fibrous ribbon, and KI fiber. The parabasal fiber provides a surface for microtubule formation, and there is one parabasal fiber for each flagellar band. The parabasal fiber possesses a dark
452:
insert into the nuclear envelope at the points of contact with the spindle poles. The nucleus maintains its characteristic position at the cell's apex through contact between kinetochores and spindle poles and apical parabasal fibers. In many other eukaryotic cells, most of the cytoplasmic
489:
has two forms: haploid and diploid. In the haploid form, it possesses two chromosomes. In the diploid form, it possesses four chromosomes. Forms with greater ploidies have also been observed, and ploidies can vary between individuals belonging to the same species of
Holomastigotoides.
443:
is located in the anterior apex of the cell, and is associated with a mitotic spindle located outside of the nucleus. This mitotic spindle is persistent throughout most of the cell cycle, which is unusual for eukaryotic cells and characteristic of
266:
lives in hindguts of lower termites, where it feeds on wood and assists the termite in wood digestion. This allows the termite to access and use nutrients found in wood that they would not have been able to digest otherwise.
892:
Jasso-Selles, Daniel E.; De
Martini, Francesca; Velenovsky, Joseph F.; Mee, Evan D.; Montoya, Samantha J.; Hileman, Jonathon T.; Garcia, Mikaela D.; Su, Nan-Yao; Chouvenc, Thomas; Gile, Gillian H. (November 2020).
406:
has been observed to change cell shape and direction of movement constantly. Intracellular calcium ion concentration affects centrin, which in turn can change flagellar band structure and basal body orientation.
415:
In the cytoplasm, food vacuoles are distributed widely and contain ingested wood. Ingested wood particles and glycogen have also been observed to be freely distributed throughout the cytoplasm.
466:
cell. The microtubules used for the cytoskeleton and mitosis are separate, and thus the cytoskeleton does not need to be disassembled for cell division to be initiated in
959:"The parabasalid symbiont community of Heterotermes aureus: Molecular and morphological characterization of four new species and reestablishment of the genus Cononympha"
834:
Gile, Gillian H.; James, Erick R.; Tai, Vera; Harper, James T.; Merrell, Trevor L.; Boscaro, Vittorio; HusnĂk, Filip; Scheffrahn, Rudolf H.; Keeling, Patrick J. (2018).
957:
Jasso-Selles, Daniel E.; De
Martini, Francesca; Freeman, Katalina D.; Garcia, Mikaela D.; Merrell, Trevor L.; Scheffrahn, Rudolf H.; Gile, Gillian H. (October 2017).
895:"The Complete Protist Symbiont Communities of Coptotermes formosanus and Coptotermes gestroi : Morphological and Molecular Characterization of Five New Species"
1296:
768:"Morphology and molecular phylogeny of Pseudotrichonympha hertwigi and Pseudotrichonympha paulistana (Trichonymphea, parabasalia) from neotropical rhinotermitids"
1309:
345:
prevent pieces of ingested wood in the termite hindgut from contacting and damaging cell surfaces. The number of flagellar bands varies based on the species of
431:
Golgi bodies can be found on the interior side of flagellar bands, spaced evenly. Endoplasmic reticulum elements can be found between Golgi and basal bodies.
335:
is the high density of flagella on the cell surface, with some reports of up to 10 000 flagella on a single cell. The organization of the flagella in
1283:
1174:
899:
840:
772:
690:
498:
have terminal centromeres. Crossing over has been observed, possibly to prevent complete segregation or no segregation of the chromatids.
1038:
836:"New Species of Spirotrichonympha from Reticulitermes and the Relationships Among Genera in Spirotrichonymphea (Parabasalia)"
505:
the same chromosomes. Chromosome division has been observed to occur in a longitudinal direction, rather than transverse. In
36:
271:
can be transferred from termite to termite by way of feeding on anal secretions of other termites during juvenile stages.
766:
Saldarriaga, Juan F.; Gile, Gillian H.; James, Erick R.; Horák, Ales; Scheffrahn, Rudolf H.; Keeling, Patrick J. (2011).
744:
147:
has notably been studied to observe the mechanisms of chromosomal pairing and segregation in haploid and diploid cells.
1257:
448:. An extranuclear matrix surrounds the nuclear envelope, except at the points where it contacts the mitotic spindle.
1314:
349:. The posterior base of Holomastigotoides cells are not flagellated, and contain vesicles that are likely used for
247:
173:, in Brazil. After initial discovery, Giovanni Battista Grassi and Anna Foa reclassified Hartmann's “male” form of
579:
strictly co-speciated with its host termites, and other mechanisms are likely involved in the phenomena observed.
226:
470:. The persistence of the extra-nuclear mitotic spindle and presence of MPM-2, a mitotic protein, indicates that
240:
219:
212:
340:
up of many basal bodies arranged in a single row, and a single flagellum emerges from each basal body, giving
205:
233:
1210:
539:
1248:
685:
78:
68:
742:
Cleveland, Lemuel R. (1960). "Pairing and segregation in haploids and diploids of
Holomastigotoides".
1128:
1350:
547:
branches separately from other genera in the
Rhinotermitidae, implying the ancestral condition of
1355:
1152:
1099:
958:
932:
865:
805:
715:
31:
533:
has been ancestrally present in this group of termites. This is supported by the observation of
1322:
1301:
1270:
1190:
1144:
1119:
1050:
1042:
978:
924:
916:
857:
797:
789:
707:
1327:
426:
cells. They are responsible for producing ATP when converting pyruvate to acetate, providing
1182:
1136:
1089:
1081:
1034:
970:
908:
849:
781:
699:
453:
microtubules are dissociated to form the mitotic spindle. However, this is not the case in
522:
295:
132:
1262:
1132:
894:
835:
703:
285:
135:
outside its nucleus during the majority of its cell cycle. As a symbiont of termites,
1344:
936:
785:
419:
394:
809:
719:
181:
in 1911, thus establishing the first use of the genus. The original host species of
1156:
1094:
869:
526:
350:
289:
281:
1275:
571:
may have speciated alongside its host termites. However, the presence of multiple
1233:
1140:
1242:
449:
393:
is a protein found in the cytoskeleton of eukaryotic cells, and plays a role in
168:
120:
974:
1186:
304:
1194:
1046:
920:
793:
711:
185:
described by
Hartmann was later invalidated due to lack of description, and
158:
was first described by Max
Hartmann in 1910. Hartmann mistakenly identified
128:
58:
48:
1148:
982:
928:
861:
801:
767:
311:
within a single host species or a result of possible co-speciation between
1054:
1022:
1227:
475:
303:
to reside in an individual host termite species. This may be a result of
1288:
1103:
1069:
390:
124:
912:
853:
1039:
10.1002/(SICI)1097-0169(1997)36:4<377::AID-CM7>3.0.CO;2-2
1204:
1117:
Cleveland, Lemuel R. (1947). "The origin and evolution of meiosis".
1085:
139:
is able to ingest wood and aid its host in digestion. In return,
1208:
1070:"The Whole Life Cycle of Chromosomes and Their Coiling Systems"
1023:"Centrin and the cytoskeleton of the protist Holomastigotoides"
278:
species have been found in multiple termite genera, including
686:"Ultrastructure of the Parabasalid Protist Holomastigotoides"
143:
is supplied with a stable habitat and steady supply of food.
127:. It is characterized by its dense, organized arrangement of
331:
is a cone-shaped cell. One of the most notable features of
575:
species in host species eliminates the possibility that
474:
spend most of their cell cycle in a suspended stage of
166:, which he observed living in a species of termite,
1217:
586:to form a monophyletic group with species found in
1074:Transactions of the American Philosophical Society
684:Lingle, Wilma L.; Salisbury, Jeffrey L. (1995).
8:
262:is an obligate symbiont of lower termites.
1205:
131:on the cell surface and the presence of a
20:
1175:"Memoirs: Observations on Trichonymphids"
1093:
543:and other genera in the Rhinotermitidae.
299:. It is possible for multiple species of
189:was subsequently named the type host for
1021:Lingle, W. L.; Salisbury, J. L. (1997).
599:
900:The Journal of Eukaryotic Microbiology
841:The Journal of Eukaryotic Microbiology
773:The Journal of Eukaryotic Microbiology
691:The Journal of Eukaryotic Microbiology
200:The following species are recognized:
162:as the female form of the parabasalid
1168:
1166:
1016:
1014:
1012:
679:
677:
675:
673:
671:
669:
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629:
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623:
7:
1010:
1008:
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1004:
1002:
1000:
998:
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994:
992:
952:
950:
948:
946:
887:
885:
883:
881:
879:
829:
827:
825:
823:
821:
819:
761:
759:
737:
735:
733:
731:
729:
621:
619:
617:
615:
613:
611:
609:
607:
605:
603:
402:direction of movement, and mitosis.
1173:Mackinnon, Doris L. (1926-04-01).
1027:Cell Motility and the Cytoskeleton
704:10.1111/j.1550-7408.1995.tb05895.x
14:
963:European Journal of Protistology
786:10.1111/j.1550-7408.2011.00575.x
35:
457:cells. The mitotic spindle of
422:, hydrogenosomes are found in
123:found in the hindgut of lower
1:
482:Chromosomes and cell division
357:Basal bodies and fiber system
193:as it is the only species of
16:Genus of flagellated protists
1141:10.1126/science.105.2724.287
582:There is strong support for
529:group, which suggests that
435:Nucleus and mitotic spindle
1372:
975:10.1016/j.ejop.2017.09.001
191:Holomastigotoides hertwigi
1187:10.1242/jcs.s2-70.278.173
1068:Cleveland, L. R. (1949).
745:Archiv fĂĽr Protistenkunde
324:Cell surface and flagella
105:
100:
32:Scientific classification
30:
23:
1181:. s2-70 (278): 173–191.
1179:Journal of Cell Science
1095:2027/coo.31924001826647
567:, which suggests that
540:Prorhinotermes simplex
557:Coptotermes testaceus
187:Coptotermes testaceus
164:Trichonympha hertwigi
1133:1947Sci...105..287C
493:The chromosomes of
255:Habitat and ecology
79:Holomastigotoididae
69:Spirotrichonymphida
197:native to Brazil.
1338:
1337:
1323:Open Tree of Life
1263:Holomastigotoides
1249:Holomastigotoides
1219:Holomastigotoides
1211:Taxon identifiers
1127:(2724): 287–289.
913:10.1111/jeu.12815
854:10.1111/jeu.12447
584:Holomastigotoides
577:Holomastigotoides
573:Holomastigotoides
569:Holomastigotoides
561:Holomastigotoides
553:Holomastigotoides
549:Holomastigotoides
535:Holomastigotoides
531:Holomastigotoides
519:Holomastigotoides
507:Holomastigotoides
502:Holomastigotoides
495:Holomastigotoides
487:Holomastigotoides
472:Holomastigotoides
468:Holomastigotoides
464:Holomastigotoides
459:Holomastigotoides
455:Holomastigotoides
446:Holomastigotoides
441:Holomastigotoides
428:Holomastigotoides
424:Holomastigotoides
404:Holomastigotoides
399:Holomastigotoides
366:Holomastigotoides
347:Holomastigotoides
342:Holomastigotoides
337:Holomastigotoides
333:Holomastigotoides
329:Holomastigotoides
313:Holomastigotoides
309:Holomastigotoides
301:Holomastigotoides
276:Holomastigotoides
274:Since discovery,
269:Holomastigotoides
264:Holomastigotoides
260:Holomastigotoides
183:Holomastigotoides
179:Holomastigotoides
160:Holomastigotoides
156:Holomastigotoides
145:Holomastigotoides
141:Holomastigotoides
137:Holomastigotoides
116:Holomastigotoides
112:
111:
96:
90:Holomastigotoides
25:Holomastigotoides
1363:
1331:
1330:
1318:
1317:
1305:
1304:
1292:
1291:
1279:
1278:
1266:
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1206:
1199:
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889:
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873:
831:
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813:
763:
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753:
739:
724:
723:
681:
559:branch with two
94:
40:
39:
21:
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1164:
1116:
1115:
1111:
1086:10.2307/1005635
1067:
1066:
1062:
1020:
1019:
990:
969:(Pt A): 48–63.
956:
955:
944:
891:
890:
877:
833:
832:
817:
765:
764:
757:
741:
740:
727:
683:
682:
601:
596:
523:Rhinotermitidae
517:The species of
515:
484:
439:The nucleus of
437:
413:
388:
379:
359:
326:
321:
315:and its hosts.
296:Anacanthotermes
257:
153:
133:mitotic spindle
93:
34:
17:
12:
11:
5:
1369:
1367:
1359:
1358:
1353:
1343:
1342:
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1280:
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1254:
1239:
1223:
1221:
1215:
1214:
1209:
1201:
1200:
1162:
1109:
1060:
1033:(4): 377–390.
988:
942:
907:(6): 626–641.
875:
848:(2): 159–169.
815:
780:(6): 487–496.
755:
725:
698:(5): 490–505.
598:
597:
595:
592:
514:
511:
483:
480:
436:
433:
412:
409:
387:
384:
378:
375:
358:
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286:Prorhinotermes
256:
253:
252:
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248:H. oxyrhynchus
244:
237:
230:
223:
216:
209:
152:
149:
119:is a genus of
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15:
13:
10:
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6:
4:
3:
2:
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803:
799:
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791:
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779:
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769:
762:
760:
756:
752:(2): 163–172.
751:
747:
746:
738:
736:
734:
732:
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726:
721:
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713:
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565:C. formosanus
562:
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528:
524:
521:found in the
520:
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395:cell division
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280:Coptotermes,
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95:Hartmann 1910
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29:
26:
22:
19:
1218:
1178:
1124:
1118:
1112:
1077:
1073:
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966:
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845:
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749:
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587:
583:
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527:monophyletic
518:
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458:
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450:Kinetochores
445:
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427:
423:
420:mitochondria
417:
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403:
398:
389:
386:Cytoskeleton
380:
371:
365:
364:
360:
351:phagocytosis
346:
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328:
327:
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308:
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294:
290:Psammotermes
282:Heterotermes
279:
275:
273:
268:
263:
259:
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246:
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232:
227:H. hartmanni
225:
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121:parabasalids
115:
114:
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106:
89:
88:
24:
18:
1243:Wikispecies
1080:(1): 1–97.
588:Coptotermes
563:species in
555:species in
418:Instead of
241:H. mirabile
220:H. bigfooti
213:H. batututi
195:Coptotermes
175:T. hertwigi
169:Coptotermes
1351:Metamonads
1345:Categories
594:References
545:P. simplex
319:Morphology
305:speciation
59:Metamonada
1356:Symbiosis
1234:Q25373728
1195:1477-9137
1047:0886-1544
937:220284193
921:1066-5234
794:1550-7408
712:1066-5234
411:Cytoplasm
377:Axostyles
353:of wood.
206:H. aureus
107:See text
49:Eukaryota
1228:Wikidata
1149:17835147
983:28942092
929:32603489
862:28710832
810:40726800
802:21895839
720:83951229
513:Taxonomy
476:prophase
234:H. minor
151:Taxonomy
129:flagella
125:termites
101:Species
75:Family:
55:Phylum:
45:Domain:
1328:1065469
1302:1120203
1289:4893330
1157:2026926
1129:Bibcode
1120:Science
1104:1005635
1055:9096959
870:3684172
525:form a
391:Centrin
85:Genus:
65:Order:
1315:104085
1193:
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935:
927:
919:
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710:
551:. Two
293:, and
1297:IRMNG
1153:S2CID
1100:JSTOR
933:S2CID
866:S2CID
806:S2CID
716:S2CID
397:. In
1310:NCBI
1284:GBIF
1276:4XZQ
1191:ISSN
1145:PMID
1051:PMID
1043:ISSN
979:PMID
925:PMID
917:ISSN
858:PMID
798:PMID
790:ISSN
708:ISSN
1271:CoL
1258:AFD
1183:doi
1137:doi
1125:105
1090:hdl
1082:doi
1035:doi
971:doi
909:doi
850:doi
782:doi
750:105
700:doi
537:in
307:of
177:to
171:sp.
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