54:
33:
312:
species. The primary feature as a plant pathogen is the intake of resources from a plant host. The initial infection occurs from asexual spores overwintered in plant debris. These spores infect the host where susceptibility is best, such as young roots of rice seedlings, or the mature sunflower head.
507:
The maintenance of the symbiosis is crucial for sporulation to occur. The endofungal bacteria possess a type III secretion system (T3SS) in order to achieve symbiosis. Mutants defective in the T3SS mechanism show reduced intracellular survival and no sporulation. This T3SS is a pathogenicity factor
382:
can be either complete sterilization, antifungal use, or the blocking of sporulation so it cannot spread. This fungal-bacterial symbiont is classified as a biosafety level 2 organism. A common method of sterilization is getting rid of all of the reproductive structures of the fungus. More difficult
398:
that allows it to communicate with its fungal host, and without the bacteria's secretion system, the fungi could not produce spores. This bacterium is passed on vertically from fungus to fungus through the sporangia while these spores are germinating. Without the bacteria none of the reproductive
287:
The initial symptoms appear as small, dispersed water-soaked spots on the back of the sunflower head. As the spots expand, mycelial growth expands into parenchyma cells, further killing cells within the head. Later stages of disease have external masses of mycelium among clumps of black sporangia,
259:
sp. The production of rhizoxin by the bacteria inhibits the ability of rice plant cells to perform mitosis, dramatically weakening or outright killing young rice seedlings. The killing of the plant cells is beneficial to both the bacteria and the fungal host, as the two live as necrotrophic
254:
use agricultural rice as a host, causing the disease Rice
Seedling Blight. This infection is first observed by the fast swelling of seedling roots, but displays no further signs of infection. The main causal agent of Rice Seedling Blight is attributed to the endosymbiotic relationship with
283:
causes the premier head rot of sunflower in South Africa. Susceptibility to disease changes throughout the age of the host. Heads inoculated at the budding stage simply do not become infected. However, when inoculated at the anthesis stage, loss was relatively high. Yield was not reduced
441:
and how the symbiosis is maintained has been an area of interest. In all cases, it is obvious that the fungus profits from the biosynthetic capabilities of the endosymbiont in order to access nutrient sources. Yet, the advantage for the bacterial symbiont is not evident.
503:
contained symbionts. Removal of the symbionts from the host degraded rhizoxin production and the symbionts were then grown in pure culture. Lastly, the re-introduction of the bacteria grown in pure culture back into the host reestablished rhizoxin production.
449:
and R. microsporus. The T3SS involved in this relationship is the first report on a T3SS involved in bacterial–fungal symbiosis. Phylogenetic analysis revealed that the T3SS represents a prototype of a clade of uncharacterized T3SSs within the
369:
causes disease in humans, one of its alternative hosts, causing infections of the lungs. In one rare case it was found tainting hospital linens in Hong Kong leading to a scare that brought the disease into the forefront of mainstream media.
797:
Jennessen, Jennifer; Nielsen, Kristian Fog; Houbraken, Jos; Lyhne, Ellen
Kirstine; SchnĂĽrer, Johan; Frisvad, Jens Christian; Samson, Robert A. (2005-02-12). "Secondary Metabolite and Mycotoxin Production by the Rhizopus microsporus Group".
426:
significantly affects sunflower yield in terms of both (oil) quality and quantity. The free fatty acid content of sunflower oil increases from 0.8% to 19.4%. Diseased sunflower plants also yielded only 81% as much seed and 55% as much oil.
483:
has developed a resistance to the toxin due to an amino acid exchange in the β-tubulin protein. The resulting necrosis of the plant tissue replenishes nutrients to both the fungus and the bacteria by feeding on the decaying matter.
406:
infection. This includes removing potential hosts not part of the system (such as wild sunflowers) that may host pests and pathogens, controlling bird feeding, and avoiding mechanical damage to the plant after its flowering.
294:
has been found to be the species involved in
Rhizopus Ear Rot of maize as well. This is characterized by small spotted sporangia structures, mycelium growth on the ear, and eventual ear and grain rot.
394:
Another way to control this pathogen would be to eliminate its bacterial endosymbiont; without this endosymbiont the fungus is unable to sporulate. The bacterium has a
854:
Fernández-PĂ©rez, RocĂo; DĂez, Lorena; González-Lázaro, Miriam; Zarazaga, Myriam; Torres, Carmen; Tenorio, Carmen; Kuipers, O. P.; Ruiz-Larrea, Fernanda (2014-05-22).
357:
This fungus is most commonly found in soil, plant debris, and foodstuffs. It is a pathogen of many crops and therefore is found in many diverse environments.
690:
Shtienberg, D. (20 Feb 2007). "Rhizopus Head Rot of
Confectionery Sunflower: Effects on Yield Quantity and Quality and Implications for Disease Management".
896:
433:
is also one of very few fungi that harbors bacterial endosymbionts to control its production of toxins. Understanding the evolutionary association between
728:
Shtienberg, D. (1997). "Rhizopus Head Rot of
Confectionery Sunflower: Effects on Yield Quantity and Quality and Implications for Disease Management".
387:
is also naturally resistant to fluconazole, ketoconazole, voriconazole and the echinocandins. Antifungal prescription drugs that usually will control
499:. Toxin formation by the bacteria has been demonstrated in analogy with Koch's postulates through the discovery that rhizoxin-producing strains of
1314:
361:
is generally found in soils with a neutral pH. These soil levels usually have lower salinity for optimum growth conditions. The growth range of
365:
ranges from 25°C to 55°C with an optimal temperature of 28°C. Its primary host is rice and it is also commonly found in maize and sunflowers.
1353:
1409:
1223:
565:
Partida-Martinez, Laila P.; Hertweck, Christian (2005-10-06). "Pathogenic fungus harbours endosymbiotic bacteria for toxin production".
1288:
164:
879:
1067:
487:
The virulence factor in all known cases are biosynthesized by the pathogenic fungus. In this case of the symbiosis between
1404:
232:
to the infected area, particularly prevalent in pre-term infants. This fungus contains the bacterial endosymbiont
1358:
475:) form a symbiotic relationship. In order to kill the living cells of its host, the harbored endobacteria secrete
1340:
471:
391:
are amphotericin B and triazoles such as posaconazole, it's also occasionally susceptible to itraconazole.
234:
211:
A domesticated variant of this species is used in the preparation of traditional soy fermentation such as
1241:
1236:
1017:
Partida-Martinez, Laila P.; Monajembashi, Shamci; Greulich, Karl-Otto; Hertweck, Christian (2007-05-01).
395:
920:
1368:
1280:
53:
288:
dispersing spores abiotically, and by birds. The diseased heads can completely rot in 3 to 7 days.
1144:
1424:
1414:
1319:
1191:
383:
sterilizations oftentimes requires control agents such as antifungals are employed. However,
321:
as a secondary cycle. The resources gathered are a result of the symbiotic relationship with
148:
574:
496:
419:
349:
reproductive cycle is replaced when put in symbiosis with the rhizoxin producing bacteria.
225:
217:
1381:
8:
1419:
836:
783:
578:
1164:
1120:
1094:"Impact of Endofungal Bacteria on Infection Biology, Food Safety, and Drug Development"
1093:
1048:
991:
958:
667:
633:"Impact of Endofungal Bacteria on Infection Biology, Food Safety, and Drug Development"
632:
606:
173:
48:
1376:
1249:
1228:
1125:
1040:
996:
978:
897:"Bedding, clothes recalled at 15 health facilities after fungus linked to two deaths"
875:
815:
745:
707:
672:
654:
598:
590:
1168:
1052:
1254:
1156:
1115:
1105:
1030:
986:
970:
867:
859:
807:
779:
737:
699:
662:
644:
610:
582:
323:
741:
703:
1110:
649:
278:
1345:
1275:
1019:"Endosymbiont-Dependent Host Reproduction Maintains Bacterial-Fungal Mutualism"
1035:
1018:
863:
1398:
1214:
982:
658:
594:
1016:
959:"Endofungal bacterium controls its host by an hrp type III secretion system"
1129:
1044:
1000:
974:
819:
764:
749:
711:
676:
602:
525:
85:
853:
1301:
871:
586:
284:
significantly when heads were inoculated at the seed development stage.
1293:
1160:
331:
318:
115:
32:
1306:
811:
1332:
1262:
1145:"Sunflower oil quality and quantity as affected by rhizopus head rot"
339:
105:
95:
65:
1185:
957:
Lackner, Gerald; Moebius, Nadine; Hertweck, Christian (2011-02-01).
856:
Industrial, medical and environmental applications of microorganisms
1327:
1267:
1208:
476:
314:
308:
268:
239:
229:
125:
1143:
Thompson, T. E.; Rogers, C. E.; Zimmerman, D. C. (1 March 1980).
495:, the hosted bacteria population produces the causative agent of
208:
is a fungal plant pathogen infecting maize, sunflower, and rice.
479:, a toxin that inhibits cell mitosis and vegetative production.
345:
There is an upcoming theory that suggests that a portion of the
327:
species, allowing for rhizoxin production to kill plant cells.
212:
75:
796:
335:
508:
that is required by the pathogen in order to cause disease.
1012:
1010:
454:
superfamily of T3SSs from plant pathogenic microorganisms.
560:
558:
556:
554:
552:
550:
548:
546:
1007:
564:
445:
Sporulation does not occur without the presence of both
1142:
543:
330:
A sexual stage is present, in the same fashion as most
276:
Head Rot in confectionery sunflower species. Alongside
956:
306:
is quite similar to the general life cycles of common
765:"Grain Molds in the Tropics: Problems and Importance"
631:Lackner, Gerald; Hertweck, Christian (2011-06-01).
422:and is a severe crop disease in Asia. In addition,
402:Preventative measures can be taken to prevent an
1396:
1091:
630:
532:. National Center for Biotechnology Information
1149:Journal of the American Oil Chemists' Society
1092:Lackner, Gerald; Hertweck, Christian (2011).
465:lives as a necrotroph where both the fungus (
313:Once infected, the production of hyphae and
317:continue to spread the infection, creating
800:Journal of Agricultural and Food Chemistry
727:
689:
31:
1119:
1109:
1068:"Managing Rhizopus Head Rot in Sunflower"
1034:
990:
666:
648:
399:structures can be created by the fungus.
1065:
762:
837:"Rhizopus Microsporus var. Microsporus"
1397:
1066:Harveson, Robert (17 September 2015).
518:
338:of alternate mating types producing a
43:sporangium containing sporangiospores
1190:
1189:
952:
950:
948:
946:
944:
942:
940:
245:
1369:3459b9ef-00a4-49d9-bbfb-fc10107c4b13
894:
831:
829:
723:
721:
626:
624:
622:
620:
1410:Fungal plant pathogens and diseases
784:10.1146/annurev.py.21.090183.001101
13:
937:
858:. Wageningen Academic Publishers.
14:
1436:
826:
718:
617:
469:) and its harbored endobacteria (
266:is similarly one of three common
238:that produces the antitumor drug
297:
52:
1136:
1085:
1059:
913:
888:
772:Annual Review of Phytopathology
457:
847:
790:
756:
683:
352:
1:
895:Zeng, Vivienne (2015-07-21).
742:10.1094/phyto.1997.87.12.1226
704:10.1094/PHYTO.1997.87.12.1226
511:
410:
373:
272:species to cause the disease
1111:10.1371/journal.ppat.1002096
650:10.1371/journal.ppat.1002096
472:Paraburkholderia rhizoxinica
235:Paraburkholderia rhizoxinica
7:
10:
1441:
1198:
1036:10.1016/j.cub.2007.03.039
864:10.3920/978-90-8686-795-0
396:type III secretion system
179:
172:
154:
147:
49:Scientific classification
47:
39:
30:
23:
528:Burkholderia rhizoxinica
1405:Fungi described in 1875
763:Williams, R.J. (1983).
975:10.1038/ismej.2010.126
925:www.life-worldwide.org
841:Joint Genome Institute
205:Rhizopus microsporus
1200:Rhizopus microsporus
901:Hong Kong Free Press
526:"Organism Overview:
497:rice seedling blight
467:Rhizopus microsporus
463:Rhizopus microsporus
431:Rhizopus microsporus
420:rice seedling blight
416:Rhizopus microsporus
292:Rhizopus microsporus
264:Rhizopus microsporus
252:Rhizopus microsporus
226:nosocomial infection
224:It can also cause a
218:Rhizopus oligosporus
191:Rhizopus microsporus
186:(Tiegh.) Mig. (1910)
158:Rhizopus microsporus
41:Rhizopus microsporus
25:Rhizopus microsporus
587:10.1038/nature03997
579:2005Natur.437..884P
250:Certain strains of
140:R. microsporus
1161:10.1007/BF02678814
378:The management of
302:The life cycle of
246:Hosts and symptoms
1392:
1391:
1377:Open Tree of Life
1192:Taxon identifiers
881:978-90-8686-243-6
812:10.1021/jf048147n
736:(12): 1226–1232.
698:(12): 1226–1232.
573:(7060): 884–888.
201:
200:
195:
187:
183:Mucor microsporus
16:Species of fungus
1432:
1385:
1384:
1372:
1371:
1362:
1361:
1349:
1348:
1346:BMSSYS0000016314
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1140:
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1123:
1113:
1089:
1083:
1082:
1080:
1078:
1063:
1057:
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1038:
1014:
1005:
1004:
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963:The ISME Journal
954:
935:
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917:
911:
910:
908:
907:
892:
886:
885:
851:
845:
844:
833:
824:
823:
806:(5): 1833–1840.
794:
788:
787:
769:
760:
754:
753:
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716:
715:
687:
681:
680:
670:
652:
628:
615:
614:
562:
541:
540:
538:
537:
522:
324:Paraburkholderia
281:, R. microsporus
257:Paraburkholderia
193:
185:
160:
57:
56:
35:
21:
20:
1440:
1439:
1435:
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1227:
1222:
1213:
1212:
1207:
1194:
1184:
1183:
1173:
1171:
1141:
1137:
1104:(6): e1002096.
1090:
1086:
1076:
1074:
1064:
1060:
1023:Current Biology
1015:
1008:
955:
938:
929:
927:
919:
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914:
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903:
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882:
852:
848:
835:
834:
827:
795:
791:
767:
761:
757:
726:
719:
688:
684:
643:(6): e1002096.
629:
618:
563:
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533:
524:
523:
519:
514:
460:
413:
376:
355:
300:
248:
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156:
143:
51:
17:
12:
11:
5:
1438:
1428:
1427:
1425:Fungus species
1422:
1417:
1415:Maize diseases
1412:
1407:
1390:
1389:
1387:
1386:
1373:
1363:
1350:
1337:
1324:
1311:
1298:
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1272:
1259:
1246:
1233:
1220:
1204:
1202:
1196:
1195:
1182:
1181:
1155:(3): 106–108.
1135:
1084:
1058:
1029:(9): 773–777.
1006:
969:(2): 252–261.
936:
912:
887:
880:
846:
825:
789:
755:
730:Phytopathology
717:
692:Phytopathology
682:
637:PLOS Pathogens
616:
542:
516:
515:
513:
510:
501:R. microsporus
493:B. rhizoxinica
489:R. microsporus
481:R. microsporus
459:
456:
447:B. rhizoxinica
439:B. rhizoxinica
435:R. microsporus
424:R. microsporus
412:
409:
404:R. microsporus
389:R. microsporus
385:R. microsporus
380:R. microsporus
375:
372:
367:R. microsporus
363:R. microsporus
359:R. microsporus
354:
351:
347:R. microsporus
304:R. microsporus
299:
296:
247:
244:
215:and sufu (see
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68:
63:
59:
58:
45:
44:
37:
36:
28:
27:
15:
9:
6:
4:
3:
2:
1437:
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1072:UNL Cropwatch
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1042:
1037:
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298:Disease cycle
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194:Tiegh. (1875)
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149:Binomial name
146:
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107:
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97:
96:Mucoromycetes
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84:
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67:
64:
61:
60:
55:
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34:
29:
26:
22:
19:
1199:
1172:. Retrieved
1152:
1148:
1138:
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1097:
1087:
1075:. Retrieved
1071:
1061:
1026:
1022:
966:
962:
928:. Retrieved
924:
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904:. Retrieved
900:
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872:10261/149561
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534:. Retrieved
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458:Pathogenesis
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86:Mucoromycota
40:
24:
18:
1302:iNaturalist
1098:PLOS Pathog
778:: 153–178.
353:Environment
332:zygomycetes
260:pathogens.
1420:Mucoraceae
1399:Categories
930:2015-10-21
906:2015-10-21
536:2014-11-18
512:References
411:Importance
374:Management
116:Mucoraceae
82:Division:
983:1751-7362
659:1553-7366
595:0028-0836
340:zygospore
319:sporangia
279:R. oryzae
134:Species:
106:Mucorales
72:Kingdom:
66:Eukaryota
1328:MycoBank
1320:10398180
1276:Fungorum
1229:60032612
1224:AusFungi
1209:Wikidata
1169:84507066
1130:21738468
1053:17827854
1045:17412585
1001:20720578
820:15740082
750:18945022
712:18945022
677:21738468
603:16208371
477:rhizoxin
315:mycelium
309:Rhizopus
274:Rhizopus
269:Rhizopus
240:rhizoxin
230:necrosis
174:Synonyms
127:Rhizopus
112:Family:
62:Domain:
1294:3386522
1215:Q309614
1121:3128126
992:3105691
668:3128126
611:4416437
575:Bibcode
418:causes
122:Genus:
102:Order:
92:Class:
1366:NZOR:
1333:177331
1307:123766
1281:177331
1268:RIZPMI
1242:566561
1167:
1128:
1118:
1051:
1043:
999:
989:
981:
921:"Life"
878:
818:
748:
710:
675:
665:
657:
609:
601:
593:
567:Nature
336:hyphae
213:tempeh
167:(1875)
165:Tiegh.
1359:58291
1315:IRMNG
1255:4SDGT
1174:7 May
1165:S2CID
1077:7 May
1049:S2CID
768:(PDF)
607:S2CID
76:Fungi
1382:9393
1354:NCBI
1289:GBIF
1263:EPPO
1237:BOLD
1176:2020
1126:PMID
1079:2020
1041:PMID
997:PMID
979:ISSN
876:ISBN
816:PMID
746:PMID
708:PMID
673:PMID
655:ISSN
599:PMID
591:ISSN
491:and
437:and
228:and
1341:NBN
1250:CoL
1157:doi
1116:PMC
1106:doi
1031:doi
987:PMC
971:doi
868:hdl
860:doi
808:doi
780:doi
738:doi
700:doi
663:PMC
645:doi
583:doi
571:437
452:hrp
221:).
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