Dog Development

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Adult dog (golden retriever)
Canine oocyte
Canine Embryo (E35-38)
Dog breeds[1]

The domestic dog (Canis lupus familiaris) has been derived from an ancestoral wolf and now consists of a breed family of more than 300 worldwide, with extensive variations in morphology (size, shape and weight). The modern dog breeds show high phenotypic diversity and are thought to have arisen from this first population bottleneck associated with wolf domestication (7,000–50,000 generations ago) and a second from more recent intensive selection to create the breed (50–100 generations ago).[2]

The average canine gestation period from ovulation to birth (parturition) is approximately 64 days and there have been identified about 400 congenital disorders relating to dog development. Many of these developmental abnormalities are common to human development.

Dog Links: Introduction | Estrous Cycle | Abnormalities | Category:Dog

Animal Development: Axolotl | Bat | Cat | Chicken | Cow | Dog | Dolphin | Echidna | Fly | Frog | Grasshopper | Guinea Pig | Hamster | Kangaroo | Koala | Lizard | Medaka | Mouse | Pig | Platypus | Rabbit | Rat | Sea Squirt | Sea Urchin | Sheep | Worm | Zebrafish | Life Cycles | Development Timetable | K12
Historic Embryology  
1897 Pig | 1900 Chicken | 1901 Lungfish | 1904 Sand Lizard | 1905 Rabbit | 1906 Deer | 1907 Tarsiers | 1908 Human | 1909 Northern Lapwing | 1909 South American and African Lungfish | 1910 Salamander | 1951 Frog | Embryology History | Historic Disclaimer

Some Recent Findings

  • Embryo biotechnology in the dog: a review[3] "Canine embryos are a scarce biological material because of difficulties in collecting in vivo-produced embryos and the inability, to date, to produce canine embryos in vitro. The procedure for the transfer of in vivo-produced embryos has not been developed adequately, with only six attempts reported in the literature that have resulted in the birth of 45 puppies. In vitro, the fertilisation rate is particularly low ( approximately 10%) and the incidence of polyspermy particularly high. So far, no puppy has been obtained from an in vitro-produced embryo. In contrast, cloning of somatic cells has been used successfully over the past 4 years, with the birth of 41 puppies reported in the literature, a yield that is comparable to that for other mammalian species. Over the same period, canine embryonic stem sells and transgenic cloned dogs have been obtained."
  • Cryopreservation of Canine Embryos[4] "Canine embryos were collected from excised reproductive organs after artificial insemination and subsequently cryopreserved by a vitrification method. When the 4-cell to morula stage of cryopreserved embryos were non-surgically transferred into the uteri of nine recipient bitches by using a cystoscope, five recipients became pregnant and four of them delivered a total of seven pups."
  • Prolonged duration of fertility of dog ova [5] "The fertile period for natural mating in dogs extends from before ovulation until day 5 post ovulation (PO) and involves a delay in oocyte maturation until 2-3 days PO and viability of secondary oocytes for 48-60 h or more. Spermatozoa do not enter the uterus after vaginal insemination in late oestrus. Cervical closure appears to occur on average 5 days PO, but conception may occur following intrauterine artificial insemination (IUAI) up to 8 days PO. Therefore, the present study was conducted to clarify the duration of fertility of canine ova. Using IUAI at 6, 7, 8 and 9 days PO (n = 5 bitches each) conception rates were 100%, 71.4%, 37.5% and 0%, respectively, with an average litter resorption rate of 30.8%, and with mean litter sizes and times to delivery PO being 4.3 +/- 1.6 and 64.3 +/- 0.3 days, 4.0 +/- 1.4 and 66.3 +/- 0.4 days, and 2.5 and 68 days for IUAI at 6, 7 and 8 days, respectively. The high pregnancy rates with IUAI at 6 and 7 days PO confirm that many canine oocytes are fertile at 4-5 days after maturation. The high rate of resorption was presumably because of aging of ova or asynchrony between embryonic development and the intrauterine environment."
More recent papers
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This table shows an automated computer PubMed search using the listed sub-heading term.

  • Therefore the list of references do not reflect any editorial selection of material based on content or relevance.
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References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.

Links: References | Discussion Page | Pubmed Most Recent | Journal Searches

Search term: Dog Embryology

Kiyoung Eun, Seon-Ung Hwang, Yeon Woo Jeong, Sunyoung Seo, Seon Yong Lee, Woo Suk Hwang, Sang-Hwan Hyun, Hyunggee Kim SV40 Large T Antigen Disrupts Embryogenesis of Canine and Porcine Somatic Cell Nuclear Transfer Embryo. Biol Proced Online: 2017, 19;13 PubMed 29075153

Lei Gao, Zhufeng Yang, Chitkale Hiremath, Susan E Zimmerman, Blake Long, Paul R Brakeman, Keith E Mostov, David M Bryant, Katherine Luby-Phelps, Denise K Marciano Afadin orients cell division to position the tubule lumen in developing renal tubules. Development: 2017, 144(19);3511-3520 PubMed 28860115

Joung Joo Kim, Kang Bae Park, Eun Ji Choi, Sang Hwan Hyun, Nam-Hyung Kim, Yeon Woo Jeong, Woo Suk Hwang Relationship between time post-ovulation and progesterone on oocyte maturation and pregnancy in canine cloning. Anim. Reprod. Sci.: 2017; PubMed 28843841

Iwona Malicka, Katarzyna Siewierska, Christopher Kobierzycki, Jedrzej Grzegrzolka, Aleksandra Piotrowska, Urszula Paslawska, Marek Cegielski, Piotr Dziegiel, Marzenna Podhorska-Okolow, Marek Wozniewski Impact of Physical Training on Sex Hormones and Their Receptors During N-Methyl-N-nitrosourea-induced Carcinogenesis in Rats. Anticancer Res.: 2017, 37(7);3581-3589 PubMed 28668849

Marta Santos, Patrícia Dias-Pereira, Carla Correia-Gomes, Ricardo Marcos, Augusto de Matos, Eduardo Rocha, Carlos Lopes Use of the optical disector in canine mammary simple and complex carcinomas. APMIS: 2017; PubMed 28586148


Dog genetics[1]
Alan Wilton (1953–2011)

NCBI Taxonomy Browser Canis lupus familiaris (Genbank common name: dog)

Synonyms: Canis familiaris, Canis domesticus, Canis canis

Chromosomes: 40 (38, X, Y)

Genetic code: Translation table 1 (Standard)

Mitochondrial genetic code: Translation table 2 (Vertebrate Mitochondrial) 16,700 bp

Lineage( full ):cellular organisms; Eukaryota; Fungi/Metazoa group; Metazoa; Eumetazoa; Bilateria; Coelomata; Deuterostomia; Chordata; Craniata; Vertebrata; Gnathostomata; Teleostomi; Euteleostomi; Sarcopterygii; Tetrapoda; Amniota; Mammalia; Theria; Eutheria; Laurasiatheria; Carnivora; Caniformia; Canidae; Canis; Canis lupus

Dog genome.jpg

Links: Dog Genome Map View | NHGRI Dog Genome Project

Dog Estrous Cycle

  • Proestrus (9 days) - Precedes estrus, estradiol concentration increases as ovarian follicules mature and the uterus enlarges. The vaginal epithelium proliferates accompanied by diapedesis of erythrocytes (most cells in vaginal smear) from uterine capillaries.
  • Estrus (9 days) - Accompanied by female mating behaviour, glandular secretions increase, the vaginal epithelium becomes hyperemic, and ovulation occurs. Cycle is influenced mainly by estrogens and the interval between successive estrus cycles is about 7 months.
  • Diestrus (70-80 days) - Accompanied by female non-mating behaviour, corpus lutea present and secretes progesterone. Uterine glands undergo hypertrophy and hyperplasia, vaginal secretions and the cervix constricts.
  • Anestrus - Anestrus is a prolonged period of sexual rest where the reproductive system is quiescent.

Oocyte Development

Transmission electron micrographs of canine geminal vesicle (GV) oocytes.[6]

Canine oocyte 04.jpg

Canine geminal vesicle (GV) oocyte.[6]

Canine oocyte 02.jpg

Dog oocyte development. (A) GV (B) GVBD (C) MI and (D) MII[6]

Oocyte to Blastocyst

Canine oocyte to blastocyst.jpg

Canine oocyte to blastocyst (Image: Dr Karine Reynaud).

Development Overview

Dog embryo at neural fold stage of development

Days shown below relate to days after ovulation (day 0).

  • 48-72 h - oocytes need to complete post-ovulatory maturation to the metaphase II stage in the isthmus of the oviduct[7]
  • 2 to 5 days - fertilization
  • 14 to 16 days - embryo attaches to uterus
  • 22 to 23 days - heartbeat visible
  • 62 to 64 days - parturition (birth or whelping)

See also Concannon 2001

Sexual differentiation begins early in the embryonic period prenatally and continues into early postnatal life.

Caudal vena cava development- five theories to origin (right-sided supracardinal, caudal cardinal, sacrocardinal, lateral sympathetic or subcardinal veins).

Carnegie Stages

Canine embryo at day 21
Canine embryo at day 40 after mating, this corresponds approximately to ~day 35-38 after fertilisation (Image: Dr Karine Reynaud).

Gestational age timed from day 0 as the day of the preovulatory serum progesterone rise in the dam, should roughly correlate with fertilisation (+/- 1 day), data[8] *calculated staging only available.

Links: Carnegie Stages

Historic Embryology

These images are from drawings by Charles Bonnet (1909), later republished in a 1921 textbook of embryology.[9]

Links: Carnegie Stage Comparison
Alaskan sled dogs, bred for their racing performance.[10]

Estrous Cycle

Estrus, also called "in heat" is the time of sexually receptivity and occurs every 17 to 21 days.

  • Ovulation occurs 5 to 6 days prior to the first day of diestrus and is indicated by plasma progesterone concentrations higher than 2 ng/mL. (Parturition (birth or whelping) occurs between 62 to 64 days after ovulation).
  • Ovulated oocytes diameter[11]
    • with the zona pellucida (167.5+/-12.7 microns)
    • without zona pellucida (133.9+/-5.3 microns)

Links: Estrous Cycle


Classified as endotheliochorial placentation forming a zonary placenta, which is a complete girdle in dogs.

Three zones:

  1. girdle zone (endotheliochorial labyrinth)
  2. hemochorial hemophagous zone (marginal hematoma)
  3. polar zone (epitheliochorial free)

Trophoblast cell invasion continues after chorioallantois villous penetration and the materno–fetal interface is described as lamellar, with fetal projections interdigitating with maternal septa.

(Data from: Miglino MA, etal., 2006[12] and other sources)

Links: Comparative Placentation - Dog

Urogenital System

Dog- male urogenital cartoon.jpg Dog- female urogenital cartoon.jpg
Male Urogenital Female Urogenital

Male Gonad

Male sex differentiation is initially mediated by Sry expression then Leydig cell produced testosterone and anti-mullerian hormone (AMH, Mis), also called mullerian-inhibiting substance (MIS) or factor (MIF).

A study using timed pregnancies and male embryo development identified testis differentiation at 36 days gestation. At this time Mullerian duct regression also commenced and was completed by 46 days gestation. Immunohistochemistry also identified Mullerian Inhibitory Substance (MIS) was present during this period in testes and was absent in the undifferentiated testis.[13]

Genital Ridge Sry and Sox9[14]

Testis induction is associated with gonadal Sry and Sox9 expression in mammals, and also with Sox9 expression in vertebrates where Sry is absent. Timing was based upon the equivalent human carnegie staging and expression was measured by quantitative reverse transcription-polymerase chain reaction (qRT-PCR).

  • Carnegie Stage 16-18 - Sry expression rose in genital ridge continuously, Sox9 expressed in both male and female genital ridge
  • Carnegie Stage 17 - Sox9 expression tenfold greater than in the ovary
  • Carnegie Stage 18 - Sry expression maximal

Chromosome 9 Sox 9

Genital Ridge Sf1 and Mis[15]

Mullerian-inhibiting substance - (Mis, Mif) Anti-mullerian hormone (AMH)

Splicing factor 1 - (Sf1) 623 amino acid protein containing a nuclear transport domain, a metal-binding or zinc finger motif, and glutamine- and proline-rich regions.

  • Carnegie Stage 15 - Sf1 expression begins in genital ridges
  • Carnegie Stage 17 - Sf1 expression pronounced in male and female gonads
  • Carnegie Stage 18 - Mis expression only in male gonads

Chromosome 20 Anti-Mullerian hormone

Links: Sry | Sox9 | Sox 9 Gene | Sf1 | Mis


The cartoons below show nanog expression in the dog during spermatogenesis.[16]

Dog- spermatozoa NANOG expression.jpg Each column represents the combination of different cell types that are present in seminiferous tubules at that specific stage.

Cell types that express NANOG are outlined in red and cell types that do not express NANOG have black and grey symbols.


  • 1–16 = steps in spermiogenesis
  • In = intermediate spermatogonia
  • B = type B spermatogonia
  • Pl = pre-leptotene stage
  • L = leptotene stage
  • Z = zygotene stage
  • P = pachytene stage
  • D = diplotene stage
  • 2nd = generation of secondary spermatocytes
  • Roman figures = stage of the epithelial cycle


Sexual Development Phases in Female of Laboratory Species[17]
Phase Rat Dog (beagle) Primate (monkey)
Neonatal Birth to postnatal Day 7 Birth–3 weeks Birth to 3–4 months
Infantile Postnatal Days 8–21 3–5 weeks Up to 29 months
Juvenile/prepubertal Postnatal Days 22–37 5 weeks–6 months Up to 43 months
Pubertal Postnatal Days 37–38 6–8 months 27–30 months

Hair Development

Coat variation in the domestic dog is governed by variants in three genes.[18]

"Coat color and type are essential characteristics of domestic dog breeds. Although the genetic basis of coat color has been well characterized, relatively little is known about the genes influencing coat growth pattern, length, and curl. We performed genome-wide association studies of more than 1000 dogs from 80 domestic breeds to identify genes associated with canine fur phenotypes. Taking advantage of both inter- and intrabreed variability, we identified distinct mutations in three genes, RSPO2, FGF5, and KRT71 (encoding R-spondin-2, fibroblast growth factor-5, and keratin-71, respectively), that together account for most coat phenotypes in purebred dogs in the United States. Thus, an array of varied and seemingly complex phenotypes can be reduced to the combinatorial effects of only a few genes."

Stem Cells

In 2009 a range of canine embryonic stem cell (ESC) lines were developed from preimplantation-stage embryos.[19]

  • maintained a normal karyotype and morphology typical of undifferentiated ESCs after multiple in vitro passages and cryopreservation.
  • embryoid bodies formed in the absence of a feeder layer in attachment or suspension culture.
  • embryoid bodies differentiated into multiple cell types.
  • ESCs introduced in vivo formed teratomas containing cell types of all three embryonic germ layers.


A recent research paper has described a new online digital atlas of the dog brain based upon anatomical and functional magnetic resonance imaging (MRI).[20]

Links: Database


Newborn bulldog with cleft palate.

There are a number of dog developmental abnormalities that are used as models for human disease.

There are currently 566 abnormality links listed on the Online Mendelian Inheritance in Animals database.

Search OMIA: Canis familiaris


  • Intersex
  • Sex reversal - not due to SRY gene translocation to an X chromosome.

Cardiac Defects

  • Canine-dilated cardiomyopathy - not associated with canine desmin.[21]

Hip dysplasia

British Veterinary Association and the German Shepherd League scoring scheme

  • scoring of nine different radiographic features of each hip
  • scale from 0 (ideal) to 6 (worst)
  • potential range of subjective scores from 0 to 108.

Links: Dog Development - Abnormalities | OMIA - Hip dysplasia


  • Congenital renal disease
  • Canine Eclampsia - (puerperal tetany, hypocalcemia) develops mainly in small-breed dogs with large litters.
  • Brucellosis - male and female can be carriers of this sexually transmitted disease.

Links: OMIA 566 abnormality links | Online Mendelian Inheritance in Animals


  1. 1.0 1.1 Heidi G Parker, Elaine A Ostrander Canine genomics and genetics: running with the pack. PLoS Genet.: 2005, 1(5);e58 PubMed 16311623 | PLoS
  2. Kerstin Lindblad-Toh, Claire M Wade, Tarjei S Mikkelsen, Elinor K Karlsson, David B Jaffe, Michael Kamal, Michele Clamp, Jean L Chang, Edward J Kulbokas, Michael C Zody, Evan Mauceli, Xiaohui Xie, Matthew Breen, Robert K Wayne, Elaine A Ostrander, Chris P Ponting, Francis Galibert, Douglas R Smith, Pieter J DeJong, Ewen Kirkness, Pablo Alvarez, Tara Biagi, William Brockman, Jonathan Butler, Chee-Wye Chin, April Cook, James Cuff, Mark J Daly, David DeCaprio, Sante Gnerre, Manfred Grabherr, Manolis Kellis, Michael Kleber, Carolyne Bardeleben, Leo Goodstadt, Andreas Heger, Christophe Hitte, Lisa Kim, Klaus-Peter Koepfli, Heidi G Parker, John P Pollinger, Stephen M J Searle, Nathan B Sutter, Rachael Thomas, Caleb Webber, Jennifer Baldwin, Adal Abebe, Amr Abouelleil, Lynne Aftuck, Mostafa Ait-Zahra, Tyler Aldredge, Nicole Allen, Peter An, Scott Anderson, Claudel Antoine, Harindra Arachchi, Ali Aslam, Laura Ayotte, Pasang Bachantsang, Andrew Barry, Tashi Bayul, Mostafa Benamara, Aaron Berlin, Daniel Bessette, Berta Blitshteyn, Toby Bloom, Jason Blye, Leonid Boguslavskiy, Claude Bonnet, Boris Boukhgalter, Adam Brown, Patrick Cahill, Nadia Calixte, Jody Camarata, Yama Cheshatsang, Jeffrey Chu, Mieke Citroen, Alville Collymore, Patrick Cooke, Tenzin Dawoe, Riza Daza, Karin Decktor, Stuart DeGray, Norbu Dhargay, Kimberly Dooley, Kathleen Dooley, Passang Dorje, Kunsang Dorjee, Lester Dorris, Noah Duffey, Alan Dupes, Osebhajajeme Egbiremolen, Richard Elong, Jill Falk, Abderrahim Farina, Susan Faro, Diallo Ferguson, Patricia Ferreira, Sheila Fisher, Mike FitzGerald, Karen Foley, Chelsea Foley, Alicia Franke, Dennis Friedrich, Diane Gage, Manuel Garber, Gary Gearin, Georgia Giannoukos, Tina Goode, Audra Goyette, Joseph Graham, Edward Grandbois, Kunsang Gyaltsen, Nabil Hafez, Daniel Hagopian, Birhane Hagos, Jennifer Hall, Claire Healy, Ryan Hegarty, Tracey Honan, Andrea Horn, Nathan Houde, Leanne Hughes, Leigh Hunnicutt, M Husby, Benjamin Jester, Charlien Jones, Asha Kamat, Ben Kanga, Cristyn Kells, Dmitry Khazanovich, Alix Chinh Kieu, Peter Kisner, Mayank Kumar, Krista Lance, Thomas Landers, Marcia Lara, William Lee, Jean-Pierre Leger, Niall Lennon, Lisa Leuper, Sarah LeVine, Jinlei Liu, Xiaohong Liu, Yeshi Lokyitsang, Tashi Lokyitsang, Annie Lui, Jan Macdonald, John Major, Richard Marabella, Kebede Maru, Charles Matthews, Susan McDonough, Teena Mehta, James Meldrim, Alexandre Melnikov, Louis Meneus, Atanas Mihalev, Tanya Mihova, Karen Miller, Rachel Mittelman, Valentine Mlenga, Leonidas Mulrain, Glen Munson, Adam Navidi, Jerome Naylor, Tuyen Nguyen, Nga Nguyen, Cindy Nguyen, Thu Nguyen, Robert Nicol, Nyima Norbu, Choe Norbu, Nathaniel Novod, Tenchoe Nyima, Peter Olandt, Barry O'Neill, Keith O'Neill, Sahal Osman, Lucien Oyono, Christopher Patti, Danielle Perrin, Pema Phunkhang, Fritz Pierre, Margaret Priest, Anthony Rachupka, Sujaa Raghuraman, Rayale Rameau, Verneda Ray, Christina Raymond, Filip Rege, Cecil Rise, Julie Rogers, Peter Rogov, Julie Sahalie, Sampath Settipalli, Theodore Sharpe, Terrance Shea, Mechele Sheehan, Ngawang Sherpa, Jianying Shi, Diana Shih, Jessie Sloan, Cherylyn Smith, Todd Sparrow, John Stalker, Nicole Stange-Thomann, Sharon Stavropoulos, Catherine Stone, Sabrina Stone, Sean Sykes, Pierre Tchuinga, Pema Tenzing, Senait Tesfaye, Dawa Thoulutsang, Yama Thoulutsang, Kerri Topham, Ira Topping, Tsamla Tsamla, Helen Vassiliev, Vijay Venkataraman, Andy Vo, Tsering Wangchuk, Tsering Wangdi, Michael Weiand, Jane Wilkinson, Adam Wilson, Shailendra Yadav, Shuli Yang, Xiaoping Yang, Geneva Young, Qing Yu, Joanne Zainoun, Lisa Zembek, Andrew Zimmer, Eric S Lander Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature: 2005, 438(7069);803-19 PubMed 16341006
  3. Sylvie Chastant-Maillard, Martine Chebrout, Sandra Thoumire, Marie Saint-Dizier, Marc Chodkiewicz, Karine Reynaud Embryo biotechnology in the dog: a review. Reprod. Fertil. Dev.: 2010, 22(7);1049-56 PubMed 20797342
  4. Yasuyuki Abe, Yoshinori Suwa, Tomoyoshi Asano, Yoshiko Yanagimoto Ueta, Nanae Kobayashi, Natsumi Ohshima, Saori Shirasuna, Mohammed Ali Abdel-Ghani, Maya Oi, Yoshiyasu Kobayashi, Masafumi Miyoshi, Kazuro Miyahara, Hiroshi Suzuki Cryopreservation of canine embryos. Biol. Reprod.: 2011, 84(2);363-8 PubMed 20926804
  5. T Tsutsui, F Takahashi, T Hori, E Kawakami, P W Concannon Prolonged duration of fertility of dog ova. Reprod. Domest. Anim.: 2009, 44 Suppl 2;230-3 PubMed 19754575
  6. 6.0 6.1 6.2 Bongkoch Turathum, Kulnasan Saikhun, Parisatcha Sangsuwan, Yindee Kitiyanant Effects of vitrification on nuclear maturation, ultrastructural changes and gene expression of canine oocytes. Reprod. Biol. Endocrinol.: 2010, 8;70 PubMed 20565987 | Reprod Biol Endocrinol.
  7. Berenice de Avila Rodrigues, José Luiz Rodrigues Influence of reproductive status on in vitro oocyte maturation in dogs. Theriogenology: 2003, 60(1);59-66 PubMed 12620580
  8. V N Meyers-Wallen Sry and Sox9 expression during canine gonadal sex determination assayed by quantitative reverse transcription-polymerase chain reaction. Mol. Reprod. Dev.: 2003, 65(4);373-81 PubMed 12840810
  9. Bailey, F.R. and Miller, A.M. (1921). Text-Book of Embryology. New York: William Wood and Co. online edition
  10. Heather J Huson, Heidi G Parker, Jonathan Runstadler, Elaine A Ostrander A genetic dissection of breed composition and performance enhancement in the Alaskan sled dog. BMC Genet.: 2010, 11;71 PubMed 20649949 | BMC Genet.
  11. H S Lee, X J Yin, Y X Jin, N H Kim, S G Cho, I H Bae, I K Kong Germinal vesicle chromatin configuration and meiotic competence is related to the oocyte source in canine. Anim. Reprod. Sci.: 2008, 103(3-4);336-47 PubMed 17212978
  12. Maria Angelica Miglino, Carlos Eduardo Ambrósio, Daniele dos Santos Martins, Cristiane Valverde Wenceslau, Christiane Pfarrer, Rudolf Leiser The carnivore pregnancy: the development of the embryo and fetal membranes. Theriogenology: 2006, 66(6-7);1699-702 PubMed 16563485
  13. V N Meyers-Wallen, T F Manganaro, T Kuroda, P W Concannon, D T MacLaughlin, P K Donahoe The critical period for mullerian duct regression in the dog embryo. Biol. Reprod.: 1991, 45(4);626-33 PubMed 1751638
  14. V N Meyers-Wallen Sry and Sox9 expression during canine gonadal sex determination assayed by quantitative reverse transcription-polymerase chain reaction. Mol. Reprod. Dev.: 2003, 65(4);373-81 PubMed 12840810
  15. Vicki N Meyers-Wallen Sf1 and Mis expression: molecular milestones in the canine sex determination pathway. Mol. Reprod. Dev.: 2005, 70(4);383-9 PubMed 15685633
  16. Ewart W Kuijk, Jeffrey de Gier, Susana M Chuva de Sousa Lopes, Ian Chambers, Ans M M van Pelt, Ben Colenbrander, Bernard A J Roelen A distinct expression pattern in mammalian testes indicates a conserved role for NANOG in spermatogenesis. PLoS ONE: 2010, 5(6);e10987 PubMed 20539761 | PLoS One.
  17. David A Beckman, Maureen Feuston Landmarks in the development of the female reproductive system. Birth Defects Res. B Dev. Reprod. Toxicol.: 2003, 68(2);137-43 PubMed 12866705
  18. Edouard Cadieu, Mark W Neff, Pascale Quignon, Kari Walsh, Kevin Chase, Heidi G Parker, Bridgett M Vonholdt, Alison Rhue, Adam Boyko, Alexandra Byers, Aaron Wong, Dana S Mosher, Abdel G Elkahloun, Tyrone C Spady, Catherine André, K Gordon Lark, Michelle Cargill, Carlos D Bustamante, Robert K Wayne, Elaine A Ostrander Coat variation in the domestic dog is governed by variants in three genes. Science: 2009, 326(5949);150-3 PubMed 19713490
  19. Andrea K Vaags, Suzana Rosic-Kablar, Cathy J Gartley, Yan Zhen Zheng, Alden Chesney, Daniel A F Villagómez, Stephen A Kruth, Margaret R Hough Derivation and characterization of canine embryonic stem cell lines with in vitro and in vivo differentiation potential. Stem Cells: 2009, 27(2);329-40 PubMed 19038794
  20. Ritobrato Datta, Jongho Lee, Jeffrey Duda, Brian B Avants, Charles H Vite, Ben Tseng, James C Gee, Gustavo D Aguirre, Geoffrey K Aguirre A digital atlas of the dog brain. PLoS ONE: 2012, 7(12);e52140 PubMed 23284904 | PMC3527386 | PLoS ONE
  21. Polona Stabej, Sandra Imholz, Serge A Versteeg, Carla Zijlstra, Arnold A Stokhof, Aleksandra Domanjko-Petric, Peter A J Leegwater, Bernard A van Oost Characterization of the canine desmin (DES) gene and evaluation as a candidate gene for dilated cardiomyopathy in the Dobermann. Gene: 2004, 340(2);241-9 PubMed 15475165


Sylvie Chastant-Maillard, Martine Chebrout, Sandra Thoumire, Marie Saint-Dizier, Marc Chodkiewicz, Karine Reynaud Embryo biotechnology in the dog: a review. Reprod. Fertil. Dev.: 2010, 22(7);1049-56 PubMed 20797342

T Poth, W Breuer, B Walter, W Hecht, W Hermanns Disorders of sex development in the dog-Adoption of a new nomenclature and reclassification of reported cases. Anim. Reprod. Sci.: 2010, 121(3-4);197-207 PubMed 20537823

Sofia Cerda-Gonzalez, Curtis W Dewey Congenital diseases of the craniocervical junction in the dog. Vet. Clin. North Am. Small Anim. Pract.: 2010, 40(1);121-41 PubMed 19942060

M J Day Immune system development in the dog and cat. J. Comp. Pathol.: 2007, 137 Suppl 1;S10-5 PubMed 17560591

Stefano Romagnoli, Donald H Schlafer Disorders of sexual differentiation in puppies and kittens: a diagnostic and clinical approach. Vet. Clin. North Am. Small Anim. Pract.: 2006, 36(3);573-606, vii PubMed 16564415


D S Martins, C E Ambrósio, N Z Saraiva, C V Wenceslau, A C Morini, I Kerkis, J M Garcia, M A Miglino Early development and putative primordial germ cells characterization in dogs. Reprod. Domest. Anim.: 2011, 46(1);e62-6 PubMed 20477984

Mohammad Shamim Hossein, Yeon Woo Jeong, Sun Woo Park, Joung Joo Kim, Eugine Lee, Kyeong Hee Ko, Park Hyuk, Song Seung Hoon, Yeun Wook Kim, Sang Hwan Hyun, Taeyoung Shin, Woo Suk Hwang Birth of Beagle dogs by somatic cell nuclear transfer. Anim. Reprod. Sci.: 2009, 114(4);404-14 PubMed 19059739

H S Lee, X J Yin, Y X Jin, N H Kim, S G Cho, I H Bae, I K Kong Germinal vesicle chromatin configuration and meiotic competence is related to the oocyte source in canine. Anim. Reprod. Sci.: 2008, 103(3-4);336-47 PubMed 17212978

M J Day Immune system development in the dog and cat. J. Comp. Pathol.: 2007, 137 Suppl 1;S10-5 PubMed 17560591

G C Luvoni, S Chigioni, M Beccaglia Embryo production in dogs: from in vitro fertilization to cloning. Reprod. Domest. Anim.: 2006, 41(4);286-90 PubMed 16869883

G D Aguirre, L F Rubin, S I Bistner Development of the canine eye. Am. J. Vet. Res.: 1972, 33(12);2399-414 PubMed 4641196

P A Holst, R D Phemister The prenatal development of the dog: preimplantation events. Biol. Reprod.: 1971, 5(2);194-206 PubMed 5165787

S Sinowatz, K H Wrobel, M F El Etreby, F Sinowatz On the ultrastructure of the canine mammary gland during pregnancy and lactation. J. Anat.: 1980, 131(Pt 2);321-32 PubMed 7462099

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Cite this page: Hill, M.A. 2017 Embryology Dog Development. Retrieved November 17, 2017, from

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© Dr Mark Hill 2017, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G