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Human Embryo stage 14 sensory placodes

Placodes are ectodermal thickenings which have important roles in development of special sensory systems.

In human development, during week 4 a series of thickened surface ectodermal patches form in pairs rostro-caudally in the head region.

Recent research suggests that all sensory placodes arise from common panplacodal ectoderm (PPE) a horseshoe-shaped region of ectoderm surrounding the anterior neural plate and neural crest. Each placode then differentiates to eventually have different developmental fates. These sensory placodes will later contribute key components of each of our special senses (vision, hearing and smell). Note that their initial postion on the developing head is significantly different to their final position in the future sensory system.

  • Adenohypophyseal placode
  • Otic placodes - the first placodes visible on the surface of the embryo.
  • Olfactory (Nasal) placodes - has 2 components (medial and lateral) and will form the nose olfactory epithelium.
  • Optic (Lens) placodes - lies on the surface, adjacent to the outpocketing of the nervous system (which will for the retina) and will form the lens.
  • Profundal/trigeminal placodes

Other species have a number of other placodes which form additional sensory structures (fish, lateral line receptor).

  • Epibranchial placodes
  • Lateral line placodes
  • Hypobranchial placodes
Other Placodes?  
Mark Hill.jpg Note that a second, later developing, form of ectodermal placode development occurs with the development of hair follicles and other integumentary specialisations. This topic does not directly relate to the specialised placodes of the head region covered here. (More? Hair Development | Integumentary_System Development)
Placode Links: Week 4 | Otic Placode | Optic Placode | Nasal Placode | Sensory System Development | Category:Placode
Historic Embryology  
1928 Sparrow Embryo | 1934 Auditory Placode

Some Recent Findings

  • Review - Transcriptional regulation of cranial sensory placode development[1]Cranial sensory placodes derive from discrete patches of the head ectoderm and give rise to numerous sensory structures. During gastrulation, a specialized "neural border zone" forms around the neural plate in response to interactions between the neural and nonneural ectoderm and signals from adjacent mesodermal and/or endodermal tissues. This zone subsequently gives rise to two distinct precursor populations of the peripheral nervous system: the neural crest and the preplacodal ectoderm (PPE). The PPE is a common field from which all cranial sensory placodes arise (adenohypophyseal, olfactory, lens, trigeminal, epibranchial, otic). Members of the Six family of transcription factors are major regulators of PPE specification, in partnership with cofactor proteins such as Eya. Six gene activity also maintains tissue boundaries between the PPE, neural crest, and epidermis by repressing genes that specify the fates of those adjacent ectodermally derived domains. As the embryo acquires anterior-posterior identity, the PPE becomes transcriptionally regionalized, and it subsequently becomes subdivided into specific placodes with distinct developmental fates in response to signaling from adjacent tissues. Each placode is characterized by a unique transcriptional program that leads to the differentiation of highly specialized cells, such as neurosecretory cells, sensory receptor cells, chemosensory neurons, peripheral glia, and supporting cells. In this review, we summarize the transcriptional and signaling factors that regulate key steps of placode development, influence subsequent sensory neuron specification, and discuss what is known about mutations in some of the essential PPE genes that underlie human congenital syndromes."
More recent papers  
Mark Hill.jpg
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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.
  • References appear in this list based upon the date of the actual page viewing.

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: Placode

Aditi Dubey, Rebecca E Rose, Drew R Jones, Jean-Pierre Saint-Jeannet Generating retinoic acid gradients by local degradation during craniofacial development: One cell's cue is another cell's poison. Genesis: 2018; PubMed 29330906

Gerhard Schlosser From so simple a beginning - what amphioxus can teach us about placode evolution. Int. J. Dev. Biol.: 2017, 61(10-11-12);633-648 PubMed 29319112

Denise A Lapa Pedreira, Gregório L Acacio, Rodrigo T Gonçalves, Renato Augusto M Sá, Reynaldo A Brandt, Ramen Chmait, Eftichia Kontopoulos, Ruben A Quintero Percutaneous fetoscopic closure of large open spina bifida using a bilaminar skin substitute. Ultrasound Obstet Gynecol: 2018; PubMed 29314321

Michael Perny, Ching-Chia Ting, Sonja Kleinlogel, Pascal Senn, Marta Roccio Generation of Otic Sensory Neurons from Mouse Embryonic Stem Cells in 3D Culture. Front Cell Neurosci: 2017, 11;409 PubMed 29311837

Raphaël Aguillon, Julie Batut, Arul Subramanian, Romain Madelaine, Pascale Dufourcq, Thomas F Schilling, Patrick Blader Cell-type heterogeneity in the early zebrafish olfactory epithelium is generated from progenitors within preplacodal ectoderm. Elife: 2018, 7; PubMed 29292696

Older papers  
  • Setting appropriate boundaries: Fate, patterning and competence at the neural plate border[2] "The neural crest and craniofacial placodes are two distinct progenitor populations that arise at the border of the vertebrate neural plate. This border region develops through a series of inductive interactions that begins before gastrulation and progressively divide embryonic ectoderm into neural and non-neural regions, followed by the emergence of neural crest and placodal progenitors. In this review, we describe how a limited repertoire of inductive signals-principally FGFs, Wnts and BMPs-set up domains of transcription factors in the border region which establish these progenitor territories by both cross-inhibitory and cross-autoregulatory interactions."
  • Graded levels of Pax2a and Pax8 regulate cell differentiation during sensory placode formation[3] "Pax gene haploinsufficiency causes a variety of congenital defects. Renal-coloboma syndrome, resulting from mutations in Pax2, is characterized by kidney hypoplasia, optic nerve malformation, and hearing loss. ..We sho.w that differential levels of zebrafish Pax2a and Pax8 modulate commitment and behavior in cells that eventually contribute to the otic vesicle and epibranchial placodes."
  • Mutual repression between Gbx2 and Otx2 in sensory placodes reveals a general mechanism for ectodermal patterning[4] "In the vertebrate head, central and peripheral components of the sensory nervous system have different embryonic origins, the neural plate and sensory placodes. This raises the question of how they develop in register to form functional sense organs and sensory circuits. Here we show that mutual repression between the homeobox transcription factors Gbx2 and Otx2 patterns the placode territory by influencing regional identity and by segregating inner ear and trigeminal progenitors. Activation of Otx2 targets is necessary for anterior olfactory, lens and trigeminal character, while Gbx2 function is required for the formation of the posterior otic placode. Thus, like in the neural plate antagonistic interaction between Otx2 and Gbx2 establishes positional information thus providing a general mechanism for rostro-caudal patterning of the ectoderm."
  • An effective assay for high cellular resolution time-lapse imaging of sensory placode formation and morphogenesis[5] "This new imaging assay provides a powerful method to analyze directly development of placode-derived sensory neurons and subsequent ganglia formation for the first time in amniotes. Viewing placode development in a head cross-section provides a vantage point from which it is possible to study comprehensive events in placode formation, from differentiation, cell ingression to ganglion assembly. Understanding how placodal neurons form may reveal a new mechanism of neurogenesis distinct from that in the central nervous system and provide new insight into how cells acquire motility from a stationary epithelial cell type."
  • Epibranchial Placodes[6] "The inner ear and the epibranchial ganglia constitute much of the sensory system in the caudal vertebrate head. ...However, recent studies indicate that both systems arise from a morphologically distinct common precursor domain: the posterior placodal area. This review summarises recent studies into the induction, morphogenesis and innervation of these systems and discusses lineage restriction and cell specification in the context of their common origin."
  • Otic Placode[7] "The inner ear epithelium, with its complex array of sensory, non-sensory, and neuronal cell types necessary for hearing and balance, is derived from a thickened patch of head ectoderm called the otic placode. ...Collectively, our results suggest that Wnt8a provides the link between FGF-induced formation of the pre-otic field and restriction of the otic placode to ectoderm adjacent to the hindbrain."
  • Postotic Placode[8] "The (zebrafish) embryonic line originates from a postotic placode that produces both a migrating sensory primordium and afferent neurons. Nothing is known about the origin and innervation of the larval lines. Here we show that a "secondary" placode can be detected at 24 h postfertilization (hpf), shortly after the primary placode has given rise to the embryonic primordium and ganglion."


Chicken Placode
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Links: Movies

Preplacodal Development

Preplacodal development model.jpg

Preplacodal development model[9]

Late Blastula Stage
  • Bmp acts as a morphogen that specifies neural crest (NC) within a narrow but low range of signalling.
  • Higher levels of Bmp signaling establish the non-neural ectoderm as a broad zone of uncommitted cells with potential to form epidermal or preplacodal ectoderm (PPE).
  • Within the non neural ectoderm
    • changing levels of Bmp do not distinguish preplacodal from epidermal potential.
    • preplacodal competence factors are uniformly induced throughout this domain.
  • expression of tfap2a/c overlaps with the lateral edges of the neural plate where, perhaps in combination with neural markers, they cell-autonomously specify NC fate.
Late Gastrula Stage (9–10 hpf)
  • preplacodal ectoderm (PPE) fate is specified in competent cells near the neural-nonneural border by dorsally expressed Bmp antagonists, Fgf and Pdgf.
  • Complete attenuation of Bmp is required for PPE specification.

Relevant markers for each ectodermal domain are shown.

Experiments carried out in zebrafish.

(Above text from figure legend[9])

Otic Placode

The otic placode is the first of the sensory placodes visible on the surface of the developing human embryo. This placode will differentiate to contribute almost entirely the components of the inner ear. The images below show the first appearance on the embryo surface during week 4 and the eventual disappearance from the surface by week 5. This is only the beginning of the complex development of this structure, influenced by the surrounding epidermis, neural tube and neural crest.

Stage 11

The scanning EM of the week 4 human embryo Carnegie stage 11 shown below is a superior dorsal view of the paired otic placodes sinking into the surface at the level of the hindbrain between day 24 and day 25.

Stage11 sem13.jpg Stage11 sem20.jpg

Stage 12

By Carnegie stage 12 26 days, only a small opening of the developing otic vesicle (otocyst) remains visible on the embryo surface located behind the second pharyngeal arch.

Stage12 sem4.jpg Stage12 sem5.jpg

Stage 13

By week 5 Carnegie stage 13 the otic vesicle (otocyst) is completely formed and is no longer visible on the embryo surface.

Stage13 sem1.jpg

Cross-sections of the embryo head at this stage show the otocyst now lies within the embryo as a hollow fluid-filled epithelial "ball", located between the epidermis and the neural tube (hindbrain).

Stage 13 serial labeled images
Stage 13 image 051.jpg Stage 13 image 052.jpg Stage 13 image 053.jpg Stage 13 image 054.jpg

Links: Inner Ear | Hearing and Balance Development

Adenohypophyseal Placode

The hypophysis, or pituitary, is an endocrine gland that links the brain to peripheral endocrine organs and systems of the body through several specific hormones. The developmental origin of the hypophysis is unique, with epithelial origins from neural ectoderm (posterior) and from surface ectoderm (anterior) the adenohypophyseal placode.

In the mouse, gonadotropin-releasing hormone-1 neurones control the release of gonadotropins from the anterior pituitary and were thought to originate from the adenohypophyseal placed. A recent study has shown that they are really associated early with the formation of the nasal placode.[10]

Drosophila and mouse placode similarity.jpg

Drosophila and mouse placode similarity[11]

Links: Pituitary Development

Olfactory Placodes


Optic Placodes

Human lens development (Carnegie stage 22, Week 8)

Optic placodes (Lens) lie on the embryo surface, adjacent to the out-pocketing of the nervous system (forms the retina) and will form the lens.

surface ectoderm -> lens placode -> lens pit -> lens vesicle -> lens fibres -> lens capsule and embryonic/fetal nucleus.

Links: Lens Development | Vision Development

Trigeminal Placodes


Links: Profundal/trigeminal placodes

Epibranchial Placodes

Zebrafish placode model[12]

Epibranchial ganglia sensory neurons formed by the facial, glossopharyngeal, and vagal placodal regions. These ganglia neurons relay from the sensory organs such as gustatory taste buds, heart baroreceptors, gut sensory enteric nerves.

Sensory System

Hearing cartoon.jpg
Senses Links: Introduction | Placodes | Hearing and Balance | Vision | Smell | Taste | Touch | Stage 22 | Category:Senses
| original Sensory page
Hearing Links: Introduction | Science Lecture | Lecture Movie | Medicine Lecture | Inner Ear | Middle Ear | Outer Ear | Balance | Hearing - Neural Pathway | Stage 22 | Abnormalities | Neonatal Diagnosis - Hearing | Hearing test | Sensory Introduction | Placodes | Student project

Categories: Hearing | Outer Ear | Middle Ear | Inner Ear

Historic Hearing Embryology 
Historic Embryology: 1880 Platypus cochlea | 1902 Development of Hearing | 1906 Membranous Labyrinth | 1910 Auditory Nerve | 1913 Tectorial Membrane | 1918 Human Embryo Otic Capsule | 1918 Cochlea | 1918 Grays Anatomy | 1922 Human Auricle | 1922 Otic Primordia | 1931 Internal Ear Scalae | 1932 Otic Capsule 1 | 1933 Otic Capsule 2 | 1936 Otic Capsule 3 | 1933 Endolymphatic Sac | 1934 Otic Vesicle | 1934 Membranous Labyrinth | 1938 Stapes - 7 to 21 weeks | 1938 Stapes - Term to Adult | 1942 Stapes - Embryo 6.7 to 50 mm | 1943 Stapes - Fetus 75 to 150 mm | 1948 Stapes - Fetus 160 mm to term | 1959 Auditory Ossicles | 1963 Human Otocyst | Historic Disclaimer

Embryo Week: Week 1 | Week 2 | Week 3 | Week 4 | Week 5 | Week 6 | Week 7 | Week 8 | Week 9

Carnegie Stages: 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | About Stages | Timeline


  1. Sally A Moody, Anthony-Samuel LaMantia Transcriptional regulation of cranial sensory placode development. Curr. Top. Dev. Biol.: 2015, 111;301-50 PubMed 25662264
  2. Andrew K Groves, Carole LaBonne Setting appropriate boundaries: fate, patterning and competence at the neural plate border. Dev. Biol.: 2014, 389(1);2-12 PubMed 24321819
  3. Matthew N McCarroll, Zachary R Lewis, Maya Deza Culbertson, Benjamin L Martin, David Kimelman, Alex V Nechiporuk Graded levels of Pax2a and Pax8 regulate cell differentiation during sensory placode formation. Development: 2012, 139(15);2740-50 PubMed 22745314
  4. Ben Steventon, Roberto Mayor, Andrea Streit Mutual repression between Gbx2 and Otx2 in sensory placodes reveals a general mechanism for ectodermal patterning. Dev. Biol.: 2012, 367(1);55-65 PubMed 22564795
  5. A Tehindrazanarivelo, H Massiou, M G Bousser [What is new in the treatment of migraine?]. [Quoi de neuf dans le traitement de la migraine?] Rev Prat: 1990, 40(5);407-10 PubMed 2155472 | BMC Neurosci.
  6. Raj K Ladher, Paul O'Neill, Jo Begbie From shared lineage to distinct functions: the development of the inner ear and epibranchial placodes. Development: 2010, 137(11);1777-85 PubMed 20460364
  7. Lisa D Urness, Christian N Paxton, Xiaofen Wang, Gary C Schoenwolf, Suzanne L Mansour FGF signaling regulates otic placode induction and refinement by controlling both ectodermal target genes and hindbrain Wnt8a. Dev. Biol.: 2010, 340(2);595-604 PubMed 20171206
  8. Andres F Sarrazin, Viviana A Nuñez, Dora Sapède, Valériane Tassin, Christine Dambly-Chaudière, Alain Ghysen Origin and early development of the posterior lateral line system of zebrafish. J. Neurosci.: 2010, 30(24);8234-44 PubMed 20554875
  9. 9.0 9.1 Hye-Joo Kwon, Neha Bhat, Elly M Sweet, Robert A Cornell, Bruce B Riley Identification of early requirements for preplacodal ectoderm and sensory organ development. PLoS Genet.: 2010, 6(9);e1001133 PubMed 20885782 | PLoS Genet.
  10. Hillery Metz, Susan Wray Use of mutant mouse lines to investigate origin of gonadotropin-releasing hormone-1 neurons: lineage independent of the adenohypophysis. Endocrinology: 2010, 151(2);766-73 PubMed 20008041
  11. Shu Wang, Natalia Tulina, Daniel L Carlin, Eric J Rulifson The origin of islet-like cells in Drosophila identifies parallels to the vertebrate endocrine axis. Proc. Natl. Acad. Sci. U.S.A.: 2007, 104(50);19873-8 PubMed 18056636 | PMC2148390
  12. Matthew N McCarroll, Alex V Nechiporuk Fgf3 and Fgf10a work in concert to promote maturation of the epibranchial placodes in zebrafish. PLoS ONE: 2013, 8(12);e85087 PubMed 24358375 | PLoS One.

Online Textbooks

Search Bookshelf placode development


Gerhard Schlosser, Cedric Patthey, Sebastian M Shimeld The evolutionary history of vertebrate cranial placodes II. Evolution of ectodermal patterning. Dev. Biol.: 2014, 389(1);98-119 PubMed 24491817

Cedric Patthey, Gerhard Schlosser, Sebastian M Shimeld The evolutionary history of vertebrate cranial placodes--I: cell type evolution. Dev. Biol.: 2014, 389(1);82-97 PubMed 24495912

Anthony Graham, Sebastian M Shimeld The origin and evolution of the ectodermal placodes. J. Anat.: 2013, 222(1);32-40 PubMed 22512454

Gerhard Schlosser Making senses development of vertebrate cranial placodes. Int Rev Cell Mol Biol: 2010, 283;129-234 PubMed 20801420

Raj K Ladher, Paul O'Neill, Jo Begbie From shared lineage to distinct functions: the development of the inner ear and epibranchial placodes. Development: 2010, 137(11);1777-85 PubMed 20460364

J Begbie, J F Brunet, J L Rubenstein, A Graham Induction of the epibranchial placodes. Development: 1999, 126(5);895-902 PubMed 9927591


Philip Barron Abitua, T Blair Gainous, Angela N Kaczmarczyk, Christopher J Winchell, Clare Hudson, Kaori Kamata, Masashi Nakagawa, Motoyuki Tsuda, Takehiro G Kusakabe, Michael Levine The pre-vertebrate origins of neurogenic placodes. Nature: 2015; PubMed 26258298

Francoise Mazet The evolution of sensory placodes. ScientificWorldJournal: 2006, 6;1841-50 PubMed 17205191

Sujata Bhattacharyya, Marianne Bronner-Fraser Hierarchy of regulatory events in sensory placode development. Curr. Opin. Genet. Dev.: 2004, 14(5);520-6 PubMed 15380243

R W Köster, R P Kühnlein, J Wittbrodt Ectopic Sox3 activity elicits sensory placode formation. Mech. Dev.: 2000, 95(1-2);175-87 PubMed 10906460

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Cite this page: Hill, M.A. 2018 Embryology Placodes. Retrieved January 19, 2018, from

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