Dog Development

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Adult dog (golden retriever)

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).[1]

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
  Historic: 1958 Early Vascular Development | 1961 Foregut Development

Animal Development: axolotl | bat | cat | chicken | cow | dog | dolphin | echidna | fly | frog | goat | grasshopper | guinea pig | hamster | horse | kangaroo | koala | lizard | medaka | mouse | opossum | pig | platypus | rabbit | rat | salamander | sea squirt | sea urchin | sheep | worm | zebrafish | life cycles | development timetable | development models | 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
Canine oocyte
Canine Embryo (E35-38)
Dog breeds[2]

Some Recent Findings

  • Assisted reproductive techniques for canines: preservation of genetic material in domestic dogs[3] "Assisted reproductive techniques (ARTs), such as artificial insemination, in vitro fertilization, and cryopreservation of gametes/zygotes, have been developed to improve breeding and reproduction of livestock and for the treatment of human infertility. Their widespread use has contributed to improvements in human health and welfare. However, in dogs, only artificial insemination using frozen semen is readily available as an ART to improve breeding and control genetic diversity. A recent priority in sperm cryopreservation is the development of alternatives to egg yolk, which is widely used as a component of the sperm extender."
  • Contrast-enhanced ultrasonography of maternal and fetal blood flows in pregnant bitches[4] "We evaluated the potential usefulness of CEUS to assess fetal-maternal circulation during pregnancy in dogs. Nine bitches were examined at 23, 30, and 45 days of gestation using an ultrasound machine (LOGIQ E9) and SonoVue® contrast media as echo-signal enhancer. Qualitative and quantitative evaluation of contrast enhancement patterns of uterine artery and utero/placental vessels were performed on recorded images. Independently of the gestational periods, the qualitative evaluation showed the initial wash-in phase from the first appearance of the uterine artery to the rapid distribution in embryonic vesicles or placenta to the progressive washout, whilst there was no enhancement of either embryos or fetuses in any bitch. Independent of gestational age, parameters derived from quantitative analysis of time intensity-curves of contrast enhancement (peak intensity, time to peak, rise time, washout) did not vary between proximal placenta, distal placenta, and uterine artery. With the progression of gestation, AUC values did not change in both proximal and distal placenta, but in the uterine artery it was lower (P ≤ 0.05) at day 30 than at day 23 (464.8 ± 16.1 vs.596.4 ± 28.1, respective" placenta
  • Computed tomographic evaluation of cleft palate in one-day-old puppies[5] "Cleft palate is a birth defect characterized by a lack of fusion between structures forming the palate. Causes include a multitude of factors, both genetic and environmental. Computed tomography (CT) is widely used to evaluate morphological features and diagnose head disorders in adult dogs. However, there is less data about its use in neonatal dogs. The purpose of this study was to perform CT evaluation of palatal defects in one-day-old puppies and to present a novel approach of 3D modeling in terms of cleft palate assessment."
More recent papers  
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More? References | Discussion Page | Journal Searches | 2019 References | 2020 References

Search term: Dog Development | Canine Development | Dog Embryology | Canine Embryology

Older papers  
These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.

See also the Discussion Page for other references listed by year and References on this current page.

  • Dynamics of male canine germ cell development[6] "Primordial germ cells (PGCs) are precursors of gametes that can generate new individuals throughout life in both males and females. Additionally, PGCs have been shown to differentiate into embryonic germ cells (EGCs) after in vitro culture. Most studies investigating germinative cells have been performed in rodents and humans but not dogs (Canis lupus familiaris). Here, we elucidated the dynamics of the expression of pluripotent (POU5F1 and NANOG), germline (DDX4, DAZL and DPPA3), and epigenetic (5mC, 5hmC, H3K27me3 and H3K9me2) markers that are important for the development of male canine germ cells during the early (22-30 days post-fertilization (dpf)), middle (35-40 dpf) and late (45-50 dpf) gestational periods. ... The PGCs were positive for POU5F1 and H3K27me3 during all assessed developmental periods, including all periods between the gonadal tissue stage and foetal testes development. The number of NANOG, DDX4, DAZL, DPPA3 and 5mC-positive cells increased along with the developing cords from 35-50 dpf. Primordial Germ Cell Development
  • Embryo biotechnology in the dog: a review[7] "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[8] "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[9] "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."


Dog genetics[2]
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.[10]

Canine geminal vesicle (GV) oocyte.[10]

Canine oocyte 02.jpg

Dog oocyte development. (A) GV (B) GVBD (C) MI and (D) MII[10]

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[11]
  • 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[12] *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.[13]

Links: Carnegie Stage Comparison
Alaskan sled dogs, bred for their racing performance.[14]

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[15]
    • 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[16] 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.[17]

Genital Ridge Sry and Sox9[12]

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[18]

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.PubmedParser error: The PubmedParser extension received invalid XML data. ()

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[19]
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.[20]</ref>

"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.[21]

  • 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.
Links: Stem Cells


A recent research paper has described a new online digital atlas of the dog brain based upon anatomical and functional magnetic resonance imaging (MRI).[22]

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.


Canine Congenital Sensorineural Deafness

Canine congenital sensorineural deafness (CCSD) is the most common cause of deafness in dogs occurring in more than 90 dog breeds.[23]

CCSD breed highest prevalence - dalmatian (most common), English setters, English cocker spaniels, bull terriers, Australian cattle dogs, whippets, catahoula leopard dogs, border collies and jack russell terriers.

This abnormality appears to be associated with intermediate cells (melanocytes, neural crest in origin) in the stria vascularis of the inner ear causing a cascade of inner ear degeneration (stria vascularis, then organ of Corti, then collapse of Reissner's membrane, finally collapse of cochlear duct.

Links: Inner Ear Melanocytes | Stria Vascularis | Inner Ear Development

Cardiac Defects

  • Canine-dilated cardiomyopathy - not associated with canine desmin.[24]

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


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Chastant-Maillard S, Chebrout M, Thoumire S, Saint-Dizier M, Chodkiewicz M & Reynaud K. (2010). Embryo biotechnology in the dog: a review. Reprod. Fertil. Dev. , 22, 1049-56. PMID: 20797342 DOI.

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