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[[File:Rat.jpg|left]]
{{Header}}
== Introduction ==
== Introduction ==
[[File:Rat.jpg|thumb]]
Rats (taxon - ''rattus'') are readily available as inbred, outbred and mutant strains. They have been generally beaten as a model by their rodent mice brethren, as the molecular tools that became available (stem cells, knockout genes, etc).


Rats (taxon- rattus) are readily available as inbred, outbred and mutant strains. They have been generally beaten as a model by their mice brethren, as the molecular tools that became available (stem cells, knockout genes, etc).


Rat embryos do have the advantage of being much larger than mouse embryos and easy to breed. Another advantage of rats is in vision development studies, as that retinal development continues postnatally (most vertebrate neurological systems are difficult to access during periods of development). Rat development is also generally 1 day behind that of mouse. (The table below gives details relating to the staging of rat development). Below this are a list of internet resources relating to the rat.


Rat embryos do have the advantage of being much larger than mouse embryos and easy to breed. Another advantage of rats is in vision development studies, as that retinal development continues postnatally (most vertebrate neurological systems are difficult to access during periods of development). Rat development is also generally 1 day behind that of mouse. (The table below gives details relating to the staging of rat development). Below this are a list of internet resources relating to the rat. This model organism has only a relatively small amount (0.2 %) of the total rat genome sequenced. (More? see rat genome)
'''Links:''' [http://embryology.med.unsw.edu.au/OtherEmb/Rat.htm original Rat page]
== Development of the Rat ==
{| class="prettytable"
| <center>'''Standard Stages'''
(Witschi)</center>
| <center>'''Age''' (days)</center>
| <center>'''Size''' (mm)</center>
| <center>'''Identification of Stages'''</center>


{{Rat}}
==Some Recent Findings==
{|
|-bgcolor="F5FAFF"
|
* '''Cross-Species Genome Wide Expression Analysis during Pluripotent Cell Determination in Mouse and Rat Preimplantation Embryos'''<ref name="PMID23077551">><pubmed>23077551</pubmed></ref> "The transition between morula and blastocyst stage during preimplantation development represents the first differentiation event of embryogenesis. Morula cells undergo the first cellular specialization and produce two well-defined populations of cells, the trophoblast and the inner cell mass (ICM).  ... This is the first study investigating the gene expression changes during the transition from morula to blastocyst in the rat preimplantation development. Our data show that in the pluripotent pool of cells of the rat and mouse preimplantation embryo substantial differential regulation of genes is present, which might explain the difficulties observed for the derivation and culture of rat ESCs using mouse conditions."
* '''Generation of germline-competent rat induced pluripotent stem cells''' <ref><pubmed>21789202</pubmed>| [http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0022008  PLoS ONE]</ref> "Our data clearly demonstrate that using only three reprogramming factors (Oct3/4, Klf4, and Sox2) rat somatic cells can be reprogrammed into a ground state. Our generated riPSCs exhibited germline transmission in either rat-rat intraspecific or mouse-rat interspecific chimeras."
* '''Comparison of staging systems for the gastrulation and early neurulation period in rodents'''<ref><pubmed>1440421</pubmed></ref>"Because there is no standard developmental staging system for the early postimplantation period of rodent embryos, investigators must now choose between a variety of systems that differ significantly. We have reviewed many of these staging systems and have summarized the ambiguities within them and the inconsistencies among them. In order to compare systems, we first obtained a consensus of the order of developmental events from the literature, and then attempted to fit existing systems into this order taking into account inconsistencies in terminology and blurred borderlines between stages."
|}
{| class="wikitable mw-collapsible mw-collapsed"
! More recent papers
|-
|-
| colspan="4" | <center>'''Cleavage and Blastula'''</center>
| [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}}


|-
Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Rat+Embryology ''Rat Embryology'']
| 1
| 1
| 0.07
| 1 cell (in oviduct)


|-
<pubmed limit=5>Rat Embryology</pubmed>
| 2
|}
| 2
==Species Stages Comparison ==
| 0.08 x 0.06
| 2 cells (in oviduct)


|-
The table below gives an approximate comparison of human, mouse and rat embryos based upon Carnegie staging.
| 3
| 3
| 0.08 x 0.05
| 4 cells (in oviduct)


|-
{{CarnegieComparisonHRM}}
| 4
| 3.5
|
| 8-12 cells (in oviduct)


|-
{{Mouse Rat Pig table}}
| 5
| 3.25
| 0.08 x 0.04
| Morula (in uterus)


|-
[[File:Rat_oocyte_01.jpg|800px]]
| 6
| 4
| (0.08 x 0.03)
| Early blastocyst (in uterus)


|-
'''Rat oocytes showing metaphase plate'''<ref><pubmed>20333307 </pubmed>| [http://www.plosone.org/article/info:doi%2F10.1371%2Fjournal.pone.0009799 PLoS One.]</ref>
| 7
| 5
| (0.12 x 0.05)
| Free blastocyst (in uterus)


|-
| colspan="4" | <center>'''Gastrula'''</center>


|-
:'''Links:''' [[Rat Timeline]]
| 8
| 6
| (0.28 x 0.07)
| Implanting blastocyst, with trophoblastic cone and inner cell mass; outgrowth of endoderm (hypoblast)


|-
==Placenta Development==
| 9
| 6.75
|
| Diplotrophoblast; inner cell mass (pendant), covered with endoderm


|-
For review of the rat placenta.<ref><pubmed>22272049</pubmed>| [http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3234607 PMC3234607] | [https://www.jstage.jst.go.jp/article/tox/24/2/24_2_95/_article }</ref>
| 10
| 7.25
| (0.3 x 0.1)
| Near complete implantation; pendant begins differentiation into embryonic and extra-embryonic parts


|-
==Adrenal and Gonad Development==
| 11
[[File:Adrenal and gonad steroidogenic factor 1 expression.jpg|600px]]
| 7.75
| (0.5 x 0.1)
| Completion of implantation; primary amniotic cyst; ectoplacental cone


|-
Adrenal and gonad steroidogenic factor 1 expression<ref><pubmed>14594453</pubmed>| [http://www.nuclear-receptor.com/content/1/1/8 Nucl Recept.]</ref>
| colspan="4" | <center>'''Primitive Streak'''</center>


|-
==References==
| 12
| 8.5
| (1.04 x 0.26)
| Connecting ectochorionic and amniotic cavities; rudiments of amniotic folds; primitive streak; start of 3rd layer formation; blastemas of heart and pericardium


|-
<references/>
| colspan="4" | <center>'''Neurula'''</center>


|-
===Reviews===
| 13
| 9
| 1.0
| Presomite neurula; fusion of chorio-amniotic folds, chorio-amniotic stalk; neural plate; embryo bent dorsally; bud of allantoic stalk


|-
===Articles===
| 14
| 9.5
| 1.5
| Somites 1-4 (occipital); pendant with 3 cavities: ectochorionic cyst, exocoelom, and amniotic cavity; ectochorionic cyst collapsing; allantoic stalk projects into exocoelom; embryo bent dorsally


|-
<pubmed>20224168</pubmed>
| 15
<pubmed>20158911</pubmed>
| 10
<pubmed>20052412</pubmed>
| 2
| Somites 5-12 (cervical); 1st visceral arch; ectochorionic cyst fused with ectoplacenta and with allantoic stalk; regression of peripheral (distal) yolk sac and trophectoderm (diplotrophoblast); Reichert's membrane; gonia in endoderm; embryo bent dorsally


|-
===Search PubMed===
| 16
| 10.5
| 2.4
| Somites 13-20 (upper thoracic); 2 visceral arches; disc and yolk sac placentas; appendicular folds; embryo reverses, curves ventrally


|-
'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=Rat+development Rat Development]
| 17
| 11
| 3.3
| Somites 21-25 (lower thoracic); yolk stalk closes at level of 15th somite; primary gonia in mesentery; primitive streak disappears; tail bud becomes organized; arm and leg buds recognizable


|-
==Additional Images==
| colspan="4" | <center>'''Tail Bud Embryo'''</center>
<gallery>
File:Reconstructed rat embryos 05.png|Reconstructed early rat embryos
File:Rat-hindbrain E11.5.jpg|Rat hindbrain E11.5, down-regulated genes and Pax6
File:Rat_thyroid_system_and_neural_development.jpg|Rat thyroid system and neural development
File:Rat neural cadherin 01.jpg
File:Rat neural cadherin 02.jpg
File:Rat neural cadherin 03.jpg
File:Rat neural cadherin 04.jpg
File:Rat neural cadherin 05.jpg
File:Rat neural cadherin 06.jpg
File:Rat neural cadherin 07.jpg
File:Rat neural cadherin 08.jpg
</gallery>


|-
==Historic Images==
| 18
<gallery>
| 11.5
File:Bailey006.jpg|Bailey Fig. 6. Schematic outline of spermatogenesis in the rat
| 3.8
File:Bailey053.jpg|Bailey Fig. 53. Four stages in the cleavage of the ovum of the white rat
| Somites 26-28 (upper lumbar); 3 visceral arches; arm buds recognizable
File:Bailey056.jpg|Bailey Fig. 56. Sections of blastocysts of the white rat, 5 days after insemination
File:Bailey061.jpg|Bailey Fig. 61. Sections of blastocysts of the white rat
File:Bailey062.jpg|Bailey Fig. 62. Cross sections of an egg-cylinder of the white rat, 8 days and 17 hours after insemination
File:Bailey063.jpg|Bailey Fig. 63 Cross sections of egg-cylinders of the white rat
File:Bailey295.jpg|Bailey Fig. 295. Ventral view of parts of the lungs, pleural cavities, peritoneal cavity, and the pleuro-peritoneal membranes in a rat embryo
</gallery>


|-
| 19
| 11.75
| 4.2
| Somites 29-31 (lower lumbar); visceral arches I-IV; cervical folds; appendicular folds and buds


|-
:'''Links:''' [[Book_-_Text-Book_of_Embryology_(1921)|Bailey, F.R. and Miller, A.M. (1921)]]
| 20
| 11.875
| 5
| Somites 32-33 (upper sacral)


|-
==External Links==
| 21
{{External Links}}
| 12
| 5.1
| Somites 34-35 (lower sacral); deep cervical sinuses


|-
* '''Rat Genome Database''' [http://rgd.mcw.edu RGD] <pubmed>17151068</pubmed>
| 22
| 12.125
| 5.2
| Somite 36 (1st caudal); olfactory pits


|-
{{Animals}}
| 23
| 12.25
| 5.6
| Somites 37-38 (caudal); start of umbilical herniation
 
|-
| 24
| 12.375
| 6
| Somites 39-40 (caudal)
 
|-
| colspan="4" | <center>'''Complete Embryo'''</center>
 
|-
| 25
| 12.5
| 6.2
| Somites 41-42 (caudal); occipital somites dispersing; 4 visceral arches; deep cervical sinuses; arm buds at somite levels 8-14, about as high as long; leg buds at somite levels 28-31, smaller; body forms a spiral of about 11/2 turns, the left face and trunk applied to yolk sac, the right side turned toward placenta; tail and allantoic stalk rise to the placenta
 
|-
| colspan="4" | <center>'''Metamorphosing Embryo'''</center>
 
|-
| 26
| 12.75
| 7
| Somites 43-45 (caudal); mandibular, maxillary, and frontonasal processes; cervical sinuses closing; mammary welts; differentiaion of handplates; arm buds vascularized, brachial nerves entering; beginning of umbilical hernia
 
|-
| 27
| 13
| 8
| Somites 46-48 (caudal); prominent facial processes and clefts; nose-snout projecting; cervical sinuses closed; primordia of mammary glands; round handplates and footplates; larger umbilical hernia
 
|-
| 28
| 13.5
| 8.5
| Somites 49-51 (caudal); 1st visceral cleft transforms into external ear duct; precartilaginous condensations in handplates
 
|-
| 29
| 14
| 9.5
| Somites 52-55 (caudal); auricular hillocks on visceral arches I and II
 
|-
| 30
| 14.5
| 10.5
| Somites 56-60 (caudal); body uncoils; mandibular precartilage; nearly round opening of external ear duct; pleuroperitoneal canal has become very narrow
 
|-
| 31
| 15
| 12
| Somites 61-63 (caudal); facial clefts closed; pleuroperitoneal canal closed; complete diaphragm
 
|-
| 32
| 15.5
| 14.2
| Somite 64 (caudal); pinna turns forward; maximal size of umbilical hernia
 
|-
| 33
| 16
| 15.5
| Somite 65 (usually this is last caudal); snout lifts off chest; last stage of metamorphosis
 
|-
| colspan="4" | <center>'''Fetus'''</center>
 
|-
| 34
| 17-18
| 16-20
 
&nbsp;
| 1st fetal stage: rapid growth of eyelids (eyes entirely covered at end of 18th day); palate complete; pinna covers ear duct; umbilical hernia withdraws
 
|-
| 35
 
ante-natal
| 19-22
| 20-40
 
&nbsp;
| 2nd fetal stage: sealed eyelids; fetal membranes and placentas reach peak of development; tail grows to 10mm;
 
'''Birth''' occurs (22nd day in rat, 19th day in mouse)
 
|-
| 35
 
post-natal
| 1-16 post-partum
| 40-1001
| After birth, fetus becomes a breathing and suckling nestling2<nowiki>; during 1st 16 days (22 to 38 days total age), eyelids remain sealed and external ear ducts plugged with periderm</nowiki>
 
|-
| 36
 
post-natal
| 17+ post-partum
| 100+1
| Periderm seals of ears and eyelids vanish; active feeding begins within next 3 days and weaning after 1 week (total weaning age, 45-48 days for rats and mice)
 
|}
'''&nbsp;'''
 
'''<nowiki>* Age (days)-</nowiki>''' days after fertilization
 
'''<nowiki>* Size (mm)-</nowiki>''' largest and smallest dimensions
 
'''Table Data Source:'''  Rat<ref>Growth. Altman and Dittmer (ed), Biological Handbooks, Fed. Soc. Exp. Biol., Washington (1962)</ref>
 
== Data For Carnegie Stages Comparison Graph (Species/Days) ==
 
{| class="prettytable"
 
|-
| '''Species'''
| '''Stage'''
| <center>'''9'''</center>
| <center>'''10'''</center>
| <center>'''11'''</center>
| <center>'''12'''</center>
| <center>'''13'''</center>
| <center>'''14'''</center>
| <center>'''15'''</center>
| <center>'''16'''</center>
| <center>'''17'''</center>
| <center>'''18'''</center>
| <center>'''19'''</center>
| <center>'''20'''</center>
| <center>'''21'''</center>
| <center>'''22'''</center>
| <center>'''23'''</center>
 
|-
| Human
| Days
| 20
| 22
| 24
| 28
| 30
| 33
| 36
| 40
| 42
| 44
| 48
| 52
| 54
| 55
| 58
 
|-
| Mouse
| Days
| 9
| 9.5
| 10
| 10.5
| 11
| 11.5
| 12
| 12.5
| 13
| 13.5
| 14
| 14.5
| 15
| 15.5
| 16
 
|-
| Rat
| Days
| 10.5
| 11
| 11.5
| 12
| 12.5
| 13
| 13.5
| 14
| 14.5
| 15
| 15.5
| 16
| 16.5
| 17
| 17.5
 
|}
 
 
'''Data Sources:''' Human<ref>Early human development and the chief sources of information on staged human embryos. O'Rahilly R. Eur J Obstet Gynecol Reprod Biol. 1979 Aug;9(4):273-80.
[http://www.ncbi.nlm.nih.gov/pubmed/400868 PMID: 400868]</ref>, Mouse<ref>The House Mouse - Atlas of Embryonic Development, Karl Theiler Springer-Verlag, NY (1972) http://genex.hgu.mrc.ac.uk/Atlas/Theiler_book_download.html</ref> and Rat.<ref>Growth. Altman and Dittmer (ed), Biological Handbooks, Fed. Soc. Exp. Biol., Washington (1962)</ref>
 
==Some Recent Findings==
 
Generation and characterization of a Tet-On (rtTA-M2) transgenic rat. Sheng Y, Lin CC, Yue J, Sukhwani M, Shuttleworth JJ, Chu T, Orwig KE. BMC Dev Biol. 2010 Feb 16;10:17. [http://www.ncbi.nlm.nih.gov/pubmed/20158911 PMID: 20158911]
 
:"This conditional expression rat model may have application for transgenic overexpression or knockdown studies of gene function in development, disease and gene therapy."
 
Sonic Hedgehog gene delivery to the rodent heart promotes angiogenesis via iNOS/netrin-1/PKC pathway. Ahmed RP, Haider KH, Shujia J, Afzal MR, Ashraf M. PLoS One. 2010 Jan 5;5(1):e8576.
[http://www.ncbi.nlm.nih.gov/pubmed/20052412 PMID: 20052412]
 
:"Reprogramming of stem cells with Shh maximizes their survival and angiogenic potential in the heart via iNOS/netrin-1/PKC signaling."
 
==References==
 
<references/>
 
'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=Rat+development Rat Development]
 
<gallery>
File:Reconstructed rat embryos.png|Reconstructed early rat embryos
File:Rat-hindbrain E11.5.jpg|Rat hindbrain E11.5, down-regulated genes and Pax6
</gallery>




{{Template:Animals}}
{{Glossary}}


{{Template:Glossary}}


{{Template:Footer}}
{{Footer}}


[[Category:Rat]]
[[Category:Rat]]

Revision as of 17:11, 18 June 2014

Embryology - 28 Mar 2024    Facebook link Pinterest link Twitter link  Expand to Translate  
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Introduction

Rat.jpg

Rats (taxon - rattus) are readily available as inbred, outbred and mutant strains. They have been generally beaten as a model by their rodent mice brethren, as the molecular tools that became available (stem cells, knockout genes, etc).


Rat embryos do have the advantage of being much larger than mouse embryos and easy to breed. Another advantage of rats is in vision development studies, as that retinal development continues postnatally (most vertebrate neurological systems are difficult to access during periods of development). Rat development is also generally 1 day behind that of mouse. (The table below gives details relating to the staging of rat development). Below this are a list of internet resources relating to the rat.


rat

Some Recent Findings

  • Cross-Species Genome Wide Expression Analysis during Pluripotent Cell Determination in Mouse and Rat Preimplantation Embryos[1] "The transition between morula and blastocyst stage during preimplantation development represents the first differentiation event of embryogenesis. Morula cells undergo the first cellular specialization and produce two well-defined populations of cells, the trophoblast and the inner cell mass (ICM). ... This is the first study investigating the gene expression changes during the transition from morula to blastocyst in the rat preimplantation development. Our data show that in the pluripotent pool of cells of the rat and mouse preimplantation embryo substantial differential regulation of genes is present, which might explain the difficulties observed for the derivation and culture of rat ESCs using mouse conditions."
  • Generation of germline-competent rat induced pluripotent stem cells [2] "Our data clearly demonstrate that using only three reprogramming factors (Oct3/4, Klf4, and Sox2) rat somatic cells can be reprogrammed into a ground state. Our generated riPSCs exhibited germline transmission in either rat-rat intraspecific or mouse-rat interspecific chimeras."
  • Comparison of staging systems for the gastrulation and early neurulation period in rodents[3]"Because there is no standard developmental staging system for the early postimplantation period of rodent embryos, investigators must now choose between a variety of systems that differ significantly. We have reviewed many of these staging systems and have summarized the ambiguities within them and the inconsistencies among them. In order to compare systems, we first obtained a consensus of the order of developmental events from the literature, and then attempted to fit existing systems into this order taking into account inconsistencies in terminology and blurred borderlines between stages."
More recent papers
Mark Hill.jpg
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More? References | Discussion Page | Journal Searches | 2019 References | 2020 References

Search term: Rat Embryology

<pubmed limit=5>Rat Embryology</pubmed>

Species Stages Comparison

The table below gives an approximate comparison of human, mouse and rat embryos based upon Carnegie staging.

Species Embryonic Comparison Timeline
Carnegie Stage
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Human Days 1 2-3 4-5 5-6 7-12 13-15 15-17 17-19 20 22 24 28 30 33 36 40 42 44 48 52 54 55 58
Mouse Days 1 2 3 E4.5 E5.0 E6.0 E7.0 E8.0 E9.0 E9.5 E10 E10.5 E11 E11.5 E12 E12.5 E13 E13.5 E14 E14.5 E15 E15.5 E16
Rat Days 1 3.5 4-5 5 6 7.5 8.5 9 10.5 11 11.5 12 12.5 13 13.5 14 14.5 15 15.5 16 16.5 17 17.5
Note these Carnegie stages are only approximate day timings for average of embryos. Links: Carnegie Stage Comparison
Table References  
Human

O'Rahilly R. (1979). Early human development and the chief sources of information on staged human embryos. Eur. J. Obstet. Gynecol. Reprod. Biol. , 9, 273-80. PMID: 400868
Otis EM and Brent R. Equivalent ages in mouse and human embryos. (1954) Anat Rec. 120(1):33-63. PMID 13207763

Mouse
Theiler K. The House Mouse: Atlas of Mouse Development (1972, 1989) Springer-Verlag, NY. Online
OTIS EM & BRENT R. (1954). Equivalent ages in mouse and human embryos. Anat. Rec. , 120, 33-63. PMID: 13207763

Rat
Witschi E. Rat Development. In: Growth Including Reproduction and Morphological Development. (1962) Altman PL. and Dittmer DS. ed. Fed. Am. Soc. Exp. Biol., Washington DC, pp. 304-314.
Pérez-Cano FJ, Franch À, Castellote C & Castell M. (2012). The suckling rat as a model for immunonutrition studies in early life. Clin. Dev. Immunol. , 2012, 537310. PMID: 22899949 DOI.


Animal Model Comparison
Postnatal Animal Models mouse rat pig
Pregnancy period (days) 18 – 21 21 – 23 110 – 118
Placenta type Discoidal, decidual
hemoendothelial choroidea
Discoidal, decidual
hemoendothelial choroidea
Epitheliochorial
Litter size 6 – 12 6 – 15 11 – 16
Birth weight (g) 0.5 – 1.5 3 – 5 900 – 1600
Weaning weight male/female (g) 18 – 25/16 – 25 55 – 90/45 – 80 6000 – 8000
Suckling period (days) 21–28 21 28–49
Solid diet beginning (days) 10 12 12 – 15
Puberty male/female (week) 4 – 6/5 6/6 – 8 20 – 28
Life expectancy (years) 1 - 2 2 - 3 14 – 18
Table data - Otis and Brent (1954)[4]   Links: timeline

Rat oocyte 01.jpg

Rat oocytes showing metaphase plate[5]


Links: Rat Timeline

Placenta Development

For review of the rat placenta.[6]

Adrenal and Gonad Development

Adrenal and gonad steroidogenic factor 1 expression.jpg

Adrenal and gonad steroidogenic factor 1 expression[7]

References

  1. ><pubmed>23077551</pubmed>
  2. <pubmed>21789202</pubmed>| PLoS ONE
  3. <pubmed>1440421</pubmed>
  4. Otis EM and Brent R. Equivalent ages in mouse and human embryos. (1954) Anat Rec. 120(1):33-63. PMID 13207763
  5. <pubmed>20333307 </pubmed>| PLoS One.
  6. <pubmed>22272049</pubmed>| PMC3234607 | [https://www.jstage.jst.go.jp/article/tox/24/2/24_2_95/_article }
  7. <pubmed>14594453</pubmed>| Nucl Recept.

Reviews

Articles

<pubmed>20224168</pubmed> <pubmed>20158911</pubmed> <pubmed>20052412</pubmed>

Search PubMed

Search Pubmed: Rat Development

Additional Images

Historic Images


Links: Bailey, F.R. and Miller, A.M. (1921)

External Links

External Links Notice - The dynamic nature of the internet may mean that some of these listed links may no longer function. If the link no longer works search the web with the link text or name. Links to any external commercial sites are provided for information purposes only and should never be considered an endorsement. UNSW Embryology is provided as an educational resource with no clinical information or commercial affiliation.

  • Rat Genome Database RGD <pubmed>17151068</pubmed>
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Cite this page: Hill, M.A. (2024, March 28) Embryology Rat Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Rat_Development

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