526:(ESCs) have the potential to allow for further understanding of the mechanisms controlling embryonic diapause. This is because the ESCs and diapausing blastocysts having very similar transcriptome profiles. ESCs are derived from the undifferentiated inner mass cells of blastocysts of an embryo – with the capability of continual proliferation in vitro. ESCs are mostly derived from mouse models, at the point where the ESCs are at optimal efficiency and are able to enter diapause.
407:, that is genetically determined, in which this process is initiated. This form of diapause is most well studied in rodents and marsupials but has been identified in many other species, including non-mammals. It is not clear how well the mechanisms studied for the onset, maintenance and release from facultative diapause in the rodent model apply to these other species.
345:
302:
gene in the fruit fly, responsible for inhibiting the formation of cyclin E-cdk2 complexes necessary for DNA synthesis. There is also evidence pointing to the upregulation of B cell translocation gene 1 (Btg1) in the mouse embryo during diapause, another known regulator of the cell cycle, responsible
294:
All multicellular organisms, from their conception, begin as a small number of cells and only grow and develop as those cells divide. In organisms which are capable of embryonic diapause, in non-ideal reproductive conditions, there is a cessation of cellular division which prevents the embryo from
285:
Experimental induction of embryonic discontinuous development within species which do not spontaneously undergo embryonic diapause in nature has been achieved; reversible developmental arrest was successfully demonstrated. This may be evidence for the evolutionary significance of this phenomenon,
319:
Specifically within placental embryonic diapause, this cessation is led by the intentional failure of the blastocyst to implant in the uterine wall, which is an essential component in developmental progression in these species. Hormones relating to the failed implantation also contribute to the
238:
Embryonic diapause is a relatively widespread phenomenon outside of mammals, with known occurrence in the reproductive cycles of many insects, nematodes, fish, and other non-mammalian vertebrates. It has been observed in approximately 130 mammalian species, which is less than two percent of all
234:
Organisms which undergo embryonic diapause are able to synchronize the birth of offspring to the most favorable conditions for reproductive success, irrespective of when mating took place. Many different factors can induce embryonic diapause, such as the time of year, temperature, lactation and
315:
has been characterized, little widely applicable characterization is available regarding entry into diapause, and the conditions which enable a blastocyst to remain dormant. Once the embryo exits diapause arrest and resumes regular development, no adverse effects are observed.
221:
Diapause provides a survival advantage to offspring, because birth or emergence of young can be timed to coincide with the most hospitable conditions, regardless of when mating occurs or length of gestation; any such gain in survival rates of progeny confers an
495:
After the vernal equinox, the photoperiod is greater than 12 hours. This decreases the production of melatonin in the pineal gland and, therefore, increases the prolactin and progesterone production in the pituitary gland and corpus luteum respectively.
418:(adj: by necessity) diapause (a.k.a. seasonal delayed implantation) is a mechanism ensuring the birth of offspring is timed during optimal environmental conditions, to ensure maximal survival. The proposed mechanism is to separate conception and
311:. Other studies have demonstrated, inversely, the lack of involvement of more common regulators of the cell cycle such as p53 within the placental model of embryonic diapause. While much of the molecular regulation involved in activating dormant
355:
With regards to the many other regulators of this form of diapause, in placental mammals, facultative diapause is most often the result of fertilization shortly following the birth of a previous litter, The consequential pups suckling during
457:, preventing continued growth of the embryo. However, in obligate diapause, the blastocyst shall enter into the dormant state in every reproductive season. This means every blastocyst a mother produces shall enter a period of diapause.
460:
Close regulation of obligate diapause is essential for survival of the mother and offspring. Premature diapause can result in forgone growth and breeding opportunities and late diapause can result in death due to adverse conditions.
425:
Obligate diapause is activated and deactivated by changes to the number of daylight hours within a day (photoperiod) and hence, occurs within specific seasons. While obligate diapause occurs in a variety of
396:
environment needed. Prolactin acting on the corpus luteum causes the progesterone level to be below optimal concentration and therefore induces embryonic facultative diapause.
352:
Facultative diapause is regulated by several factors, including the maternal environment and ovarian competency, the pituitary gland, and metabolic stress and lactation.
295:
growing and maturing, delaying the maturation of the embryo until conditions are ideal enough to promote the survival of the offspring, and in some cases, the mother.
541:
pathways. These similarities allow for the potential to use ESCs as a cellular model to identify the molecular factors which regulate embryonic diapause.
37:
136:
336:
occurs regularly in the reproductive cycle of the affected species, and is often associated with seasonal changes and photo-period.
332:
occurs in response to certain environmental or metabolic stressors, such as drastic changes in temperature, feeding, or lactation.
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61:
515:
environment. The presence of putrescine may indicate a role in inducing the escape of the embryo from obligate diapause.
986:
Murphy BD (December 2012). "Embryonic diapause: Advances in understanding the enigma of seasonal delayed implantation".
286:
with latent capacity for diapause potentially present in a much wider segment of species than known to occur naturally.
827:"Embryonic developmental arrest in the annual killifish Austrolebias charrua: A proteomic approach to diapause III"
1038:
Deng L, Li C, Chen L, Liu Y, Hou R, Zhou X (November 2018). "Research advances on embryonic diapause in mammals".
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195:
100:
500:
529:
Both diapausing blastocysts and ESCs have transcriptome profile similarities, including downregulation of
129:
56:
431:
214:
until conditions allow for attachment to the uterine wall to proceed as normal. As a result, the normal
169:
110:
618:
933:
404:
328:
There are two distinct forms of embryonic diapause, characterized by different conditions of onset.
594:
586:
523:
203:
1194:"Transcriptional profiling of lung cell populations in idiopathic pulmonary arterial hypertension"
1231:
1063:
769:
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619:"Review: Embryonic diapause in the European roe deer – slowed, but not stopped - ScienceDirect"
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in mammals) is a reproductive strategy used by a number of animal species across different
538:
503:). The Odc gene produces the ODC protein, a rate-limiting enzyme in the production of the
481:
276:
157:
825:
Chalar C, Clivio G, Montagne J, Costábile A, Lima A, Papa NG, Berois N, Arezo MJ (2021).
937:
298:
Regulation of the cell cycle as it relates to embryonic diapause has been linked to the
1218:
1193:
1192:
Saygin D, Tabib T, Bittar HE, Valenzi E, Sembrat J, Chan SY, et al. (April 2020).
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1272:
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803:
582:
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271:). Some groups only have one species that undergoes embryonic diapause, such as the
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492:, preventing development of the blastocyst. This induces embryonic diapause.
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477:
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1134:"Molecular Regulators of Embryonic Diapause and Cancer Diapause-like State"
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1059:
1007:
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Ptak GE, Tacconi E, Czernik M, Toschi P, Modlinski JA, Loi P (2012-03-12).
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Similarly to facultative diapause, a series of hormonal changes arrest the
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468:, the photoperiod is less than 12 hours. This increases the production of
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in different groups, there are significant variations in diapause length.
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83:
78:
28:
1132:
Hussein AM, Balachandar N, Mathieu J, Ruohola-Baker H (September 2022).
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640:
172:, and is characterized by a dramatic reduction or complete cessation of
738:
Desmarais JA, Bordignon V, Lopes FL, Smith LC, Murphy BD (March 2004).
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In placental embryonic diapause, the blastocyst does not immediately
161:
46:
480:, this increase in melatonin decreases prolactin secretion from the
422:(birth) so that each can occur at the most favourable time of year.
344:
572:, from late 19th century English: dia- 'through' + pause- 'delay'.
381:
343:
244:
499:
The increase in prolactin induces expression of the gene Odc (
403:
that undergoes facultative diapause tends to have a specific
476:. Due to the inhibitory relationship between melatonin and
1254:
915:
913:
388:. The main function of the corpus luteum is to secrete
372:
in a pregnant female. The corpus luteum is a temporary
740:"The escape of the mink embryo from obligate diapause"
639:Lopes FL, Desmarais JA, Murphy BD (December 2004).
484:. The decrease in prolactin consequently decreases
210:, but rather remains in this non-dividing state of
1083:"Trophoblast-uterine interactions at implantation"
882:The International Journal of Developmental Biology
164:where this takes place, the process occurs at the
922:"Embryonic diapause is conserved across mammals"
446:) only have a diapause of around fourteen days.
376:that is formed from the leftover cells from the
218:period is extended for a species-specific time.
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790:Renfree MB, Shaw G (March 2000). "Diapause".
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16:Evolutionary reproductive strategy of mammals
8:
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1021:
1019:
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348:Mechanism of facultative embryonic diapause
438:) have a diapause of around 200 days while
876:Fenelon JC, Banerjee A, Murphy BD (2014).
392:during pregnancy in order to maintain the
239:species of mammals. These include certain
137:
123:
18:
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1108:
1098:
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878:"Embryonic diapause: Development on hold"
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842:
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689:Renfree MB, Fenelon JC (September 2017).
656:
641:"Embryonic diapause and its regulation"
610:
562:
26:
1087:Reproductive Biology and Endocrinology
364:to be released. This in turn reduces
7:
691:"The enigma of embryonic diapause"
14:
1081:Aplin JD, Kimber SJ (July 2004).
1052:10.1016/j.anireprosci.2018.09.009
988:Reproduction in Domestic Animals
804:10.1146/annurev.physiol.62.1.353
384:, once it has released a mature
303:for inhibiting transition from G
35:
176:, arresting most often in the
1:
757:10.1095/biolreprod.103.023572
947:10.1371/journal.pone.0033027
844:10.1371/journal.pone.0251820
160:. In more than 130 types of
1040:Animal Reproduction Science
792:Annual Review of Physiology
1295:
581:The vernal equinox is the
1255:"Roe Deer Research Group"
994:(Supplement 6): 121–124.
230:Evolutionary significance
101:Critical thermal maximum
744:Biology of Reproduction
501:ornithine decarboxylase
1100:10.1186/1477-7827-2-48
453:development, prior to
432:Western spotted skunks
349:
1279:Developmental biology
1198:Pulmonary Circulation
1151:10.3390/cells11192929
895:10.1387/ijdb.140074bm
551:Weddell seal#Breeding
347:
170:embryonic development
111:Developmental biology
524:Embryonic stem cells
519:Embryonic stem cells
464:Prior to the vernal
340:Facultative diapause
330:Facultative diapause
206:has resulted in the
154:delayed implantation
938:2012PLoSO...733027P
658:10.1530/rep.1.00444
595:southern hemisphere
587:northern hemisphere
405:developmental stage
368:secretion from the
204:sexual reproduction
22:Part of a series on
708:10.1242/dev.148213
488:production in the
436:Spilogale gracilis
350:
320:embryonic arrest.
158:biological classes
150:Embryonic diapause
89:Embryonic diapause
1210:10.1111/een.12792
1000:10.1111/rda.12046
701:(18): 3199–3210.
591:September equinox
411:Obligate diapause
334:Obligate diapause
290:General mechanism
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1247:Further reading
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1258:. Retrieved
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1144:(19): 2929.
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645:Reproduction
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535:biosynthesis
528:
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486:progesterone
474:pineal gland
463:
459:
455:implantation
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390:progesterone
366:progesterone
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280:Artiodactyla
237:
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224:evolutionary
220:
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153:
149:
148:
88:
74:Cryptobiosis
57:Hibernaculum
695:Development
420:parturition
313:blastocysts
226:advantage.
96:Winter rest
69:Aestivation
52:Hibernation
1260:2008-04-05
605:References
589:, and the
531:metabolism
509:putrescine
451:blastocyst
265:marsupials
249:armadillos
166:blastocyst
1236:202006476
1093:(1): 48.
505:polyamine
478:prolactin
470:melatonin
362:prolactin
360:promotes
358:lactation
269:kangaroos
253:mustelids
216:gestation
168:stage of
79:Brumation
1273:Category
1228:32166015
1170:36230891
1119:15236654
1068:52884156
1060:30266523
1046:: 1–10.
1008:23279480
966:22427933
926:PLOS ONE
904:25023682
863:34086690
831:PLOS ONE
812:10845095
774:38759201
766:14585805
717:28928280
667:15579584
570:Diapause
545:See also
416:Obligate
273:roe deer
212:dormancy
84:Diapause
29:dormancy
1219:7052475
1161:9562880
957:3299720
934:Bibcode
854:8177498
725:6441064
593:in the
585:in the
513:uterine
472:in the
466:equinox
428:species
401:species
394:uterine
380:in the
275:in the
263:), and
261:badgers
257:weasels
241:rodents
198:in the
196:implant
162:mammals
27:Animal
1234:
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772:
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665:
300:dacapo
267:(e.g.
255:(e.g.
208:zygote
202:after
200:uterus
47:Torpor
1232:S2CID
1138:Cells
1064:S2CID
770:S2CID
721:S2CID
557:Notes
399:Each
382:ovary
324:Types
277:order
245:bears
189:phase
106:Sleep
1224:PMID
1166:PMID
1115:PMID
1056:PMID
1004:PMID
962:PMID
900:PMID
859:PMID
808:PMID
762:PMID
713:PMID
663:PMID
537:and
386:ovum
259:and
1214:PMC
1206:doi
1156:PMC
1146:doi
1105:PMC
1095:doi
1048:doi
1044:198
996:doi
952:PMC
942:doi
890:doi
849:PMC
839:doi
800:doi
752:doi
703:doi
699:144
653:doi
649:128
183:or
62:HIT
1275::
1230:.
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1212:.
1202:10
1200:.
1196:.
1178:^
1164:.
1154:.
1142:11
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1136:.
1113:.
1103:.
1089:.
1085:.
1062:.
1054:.
1042:.
1016:^
1002:.
992:47
990:.
974:^
960:.
950:.
940:.
928:.
924:.
912:^
898:.
886:58
884:.
880:.
857:.
847:.
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