Carnegie stage 17

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Introduction

Facts

Week 6, 42 - 44 days, 11 - 14 mm

Gestational Age GA week 8

Summary

Ectoderm: sensory placodes, lens pit, otocyst,nasal pits moved ventrally, fourth ventricle of brain
Mesoderm: heart prominence
Head: 1st, 2nd and 3rd pharyngeal arch, forebrain, eye, auricular hillocks
Body: heart, liver, umbilical cord, mesonephric ridge
Limb: upper and lower limb buds, hand digital rays

Features

pigmented eye, nasal pit, nasolacrimal groove, external acoustic meatus, auricular hillock, heart, digital rays, liver pronminance, thigh, ankle, foot plate, umbilical cord
Identify: pigmented eye, nasal pit, nasolacrimal groove, external acoustic meatus, auricular hillock, heart, digital rays, liver prominence, thigh, ankle, foot plate, umbilical cord

Historic Papers: 1926 Precervical Sinus | 1948 stage 15-18

Week:	1	2	3	4	5	6	7	8
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

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

Kyoto Collection

View: This is a left lateral view of embryo. Amniotic membrane removed.

Ventral view of head region (1 mm scale).

Human Embryo (Carnegie stage 17)
Embryo and membranes
Embryo unlabeled
Embryo CRL
Embryo features
Membranes
Membrane measurements
Oral cavity roof unlabeled
Oral cavity roof features

Right lateral view of embryo enclosed in chorionic sac. scale bar 5 mm.

Kyoto embryo (16834) showing detail of umbilicus Carnegie stage 17 (1 mm scale bar)

‎‎MRI Stage 17

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This is a MRI off-axis sagittal (not in at the exact anatomical plane) section through the week 6 embryo

Image source: The Kyoto Collection images are reproduced with the permission of Prof. Kohei Shiota and Prof. Shigehito Yamada, Anatomy and Developmental Biology, Kyoto University Graduate School of Medicine, Kyoto, Japan for educational purposes only and cannot be reproduced electronically or in writing without permission.

Carnegie Collection

Carnegie Collection Embryos - Stage 17
Serial No.	Size (mm)	Grade	Fixative	Embedding Medium	Plane	Thinness (µm)	Stain	Year	Notes
353	E, 11.0 Ch., 40x35x20	Good	Formalin	P	Coronal	10	(Stain - Haematoxylin Eosin)	1906	Very advanced
485	E, 13.0 Ch., 33x25	Exc.	Formalin	P	Coronal	40	Al. coch.	1911	Injected (India ink)
544	E., 11.5 Ch., 30	Good	Zenker-Formol	P	Sagittal	40	Al. coch.	1911	Operative Injected (India ink)
562	E, 13.0 Ch., 28x17x17	Poor	Formalin	P	Sagittal	100	Al. coch.	1912	Advanced
623	E 10.1	Good	Alc.	P	Transverse	20	H. & Congo red	1912	Operative. Median in group
695	E, 13.5 Ch., 40x40x17	Poor	Formalin	P	Transverse	10	H. & Congo red	1913	Macerated
916	E, 11.0 Ch, 30x30x16	Good	Bouin	C	Transverse	40	H.&E, or. G.	1915	Most-advanced third
940	E, 14.0 Ch, 28x23x21	Good	Formalin	C	Transverse	40	H.&E, or. G.	1914	Advanced
1232	E, 14.5 Ch., 35x35x30	Poor	Formalin	P	Coronal	40	Al. coch.	1915	Close to No.1267A
1267A	E-a 145 Ch., 35x30x26 40	Good	Formalin	C	Sagittal	20	(Stain - Haematoxylin Eosin). or. G	1915	Excellent CN.S.
1267B ??	E, 125	Good	Formalin	p	Sagittal	20	(Stain - Haematoxylin Eosin)	1917	Tubal
5642	E, 11.5 Ch.. 33x30x17	Good	Formalin	p	Transverse	15	Al. coch.	1928	Right upper limb injured
5893	E.. 13.2	Good	Formalin	C-P	Transverse	20	Al. coch.	1929	Most advanced in group
6258	E. 14.0 Ch 48x35x25	Good	Formalin	C-P	Transverse	10	(Stain - Haematoxylin Eosin)	1930	Median in group
6519	E.. 10.8*	Exc	Corrov acetic	C-P	Sagittal	8	Al. coch.	?	Least-advanced or middle third
6520	E., 14.2*	Exc.	Corros. acetic	C-P	Transverse	10	Al. coch.	P	Median in group. Ag added to slides 1-25
6521	E., 13.2*	Exc.	Corros. acetic	C-P	Transverse	8-18	Al. coch.	?	Sections vary in thinness
6631	E., 13.0	Good	Formalin	C-P	Coronal	10	(Stain - Haematoxylin Eosin)	1932	Tubal. Advanced
6742	E, 11.0 Ch.,50x40x15	Good	Formalin	C-P	Transverse	12	H. & phlox.	1933	Good primary germ cells
6758	E., 12.8	Good	Formalin	C-P	Transverse	10	H. & phlox.	1933	Least—advanced third
7317	E, 10.0*	Good		P	Coronal	10	(Stain - Haematoxylin Eosin)	1936	His embryo “Ru." Every third section
7436	E., 13.0	Good	Formalin	C-P	Coronal	30	Al. coch.	1937	Most-advanced third
8101	E., 13.0	Exc.	Bouin	C-P	Transverse	10	(Stain - Haematoxylin Eosin)	1943	Operative
8118	E., 12.6	Exc.	Bouin	C-P	Coronal	10	(Stain - Haematoxylin Eosin)	1943	Middle third
8253	E., 11.2 Ch.,30x20x10	Good	Bouin	C-P	Coronal	10	Al. coch., phlox.	1944	Operative. Least advanced in group
8789	E., 11.7	Exc.	Bouin	C-P	Sagittal	10	Azan	1950
8969	E., 11.2	Exc.	?	p	Transverse	15	Azan	1919	Univ. Chicago No. H566
8998	E., 11.0	Exc.	?	C-P	Coronal	10	Azan	1952
9100	E., 12.0 Ch., 12x13x10	Exc.	Formol-chrom. subl.	C-P	Sagittal	10	Azan	1933	Univ. Chicago No. H1475
9282	E, 12.0 Ch., 16	Good	Ale.	p	Transverse	15	Ag	1955	Mislaid
Abbreviations Size - E. is the greatest length of the embryo and Ch. is the mean diameter of the chorion. Grade - total grade of the specimen and includes both its original quality and the condition of the mounted sections. Embedding medium - paraffin (P) or a combination of celloidin and paraffin (C-P). Fixative - formalin (Formol), alcohol and formalin (Alc, formol), Bouin (Bouin solution) Stain - ? - unknown or not determined.

iBook - Carnegie Embryos
	iTunes link \| iBook Store Description - Imagine the excitement of seeing this incredible early period of human development for the first time. Now consider that much of our initial understanding of human development is based upon study of historic embryo collections. You can now look at these historic images of the first 8 weeks after fertilisation and explore for yourself the changes that occur in human development during this key period. This current book is designed as an atlas of the Carnegie embryo stages with some brief notes and additional information covering the first 8 weeks of development. These images are from from the beginning of last century and are one of the earliest documented series of human embryos collected for basic research and medical education on development. I hope you enjoy learning about the amazing early events that begin to make and shape us. This is the second book in a series of educational releases from UNSW Embryology. Release: First Edition - Mar 12, 2012 ISBN 978-0-7334-3148-7 Print Length 82 Pages, 25.8 MB Language English. PDF Preview version 3.87 MB (Read the associated information, this is an edited educational preview version with many features not functioning). The current website also includes numerous embryo images from this textbook (see Embryonic Development and Carnegie Embryos).

Eye, muscle primordia, and associated nerves
Embryo 6258 Eye muscle primordia
Embryo 6258 Brain, eye, principal nerves
Embryo 6258 Dorsocranial aspect
Embryo 6258 Ventrocaudal aspect

Embryo 544
Embryo 544
Embryo 544
Embryo 544

Blechschmidt Collection

‎‎Embryo 10mm

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Model from serial section reconstruction.

Model Embryo 10mm Movie
model left
model ventral
model right
model dorsal
model cranial end
model placental cord

Image source: The Blechschmidt Collection images are reproduced with the permission of Prof. Christoph Viebahn, director of the Institute of Anatomy and Embryology, , University Medical Center Göttingen. Images are for educational purposes only and cannot be reproduced electronically or in writing without permission.

Hill Collection

Hill H58

left dorsolateral	left lateral
Hill H202

Embryo Virtual Slide

Stage 17 - Left Lateral

‎‎Mobile | Desktop | Original

Stage 17 | Embryo Slides

Stage 17 - Ventral View

‎‎Mobile | Desktop | Original

Stage 17 | Embryo Slides

Image source: The images from the Hill Collection (part of the Embryological Collection) are reproduced with the permission of the Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity. Images are for educational purposes only and must not be reproduced electronically or in writing without permission from the Museum für Naturkunde Berlin.

Hinrichsen Collection

Hinrichsen collection Human Embryo ME16 (stage17).

Note the developing mandible and maxilla in this ventrolateral view of the head.

The developing maxilla in this ventral view of the nasal opening.

Image source: The Hinrichsen Collection images are reproduced with the permission of Prof. Beate Brand-Saberi, Head, Department of Anatomy and Molecular Embryology, Ruhr-Universität Bochum. Images are for educational purposes only and cannot be reproduced electronically or in writing without permission.

Madrid Collection

Madrid Collection Embryos
Carnegie Stage	Embryo	Days	CRL (mm)	Section thickness	Staining	Section plane
17	CN 4	41	11	10	(Stain - Haematoxylin Eosin)	transverse
17	J1	41	11.5	10	Bielschowsky (Stain - Haematoxylin Eosin)	frontal
17	MAR 4	42	12	10	(Stain - Haematoxylin Eosin)	transverse
17	VE-4	43	12.5	10	(Stain - Haematoxylin Eosin)	transverse

Scanning EM


Ventral view of head showing upper lip, maxilla and nasal region.	Note that a ventral image of only half the head has been "mirrored" to generate this image. Image Source: Prof Virginia Diewert

Movies

MRI

‎‎MRI Stage 17

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Tomography

‎‎Stage 17 Embryo

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Optical projection tomography movie of rotating stage 17 embryo.

Note the detailed structural view of neural system development.

Stage 17 Optical Projection Tomography (left)

Stage 17 Optical Projection Tomography (right)

Links: Movies

Other Specimens

Volcher R. Le systeme nerveux pe'riphe'rique cninien d'un embryon humain de 12 mm. (1959) Arch. Biol. (Liege), 70:179-215. describes the peripheral nervous system.

Hendrix MJ Brailey JL and Shenker L. SEM-dissection of a human embryo derived from an ectopic pregnancy. (1985) Early Hum Dev. 11(1): 61-8. PMID 4006825

Events

neural
- telencephalon areas of the future archicortex, paleocortex, and neocortex, visible. Beginning of future choroid plexus^[1]
- primordium of the epidural space appears first on the ventral part of the vertebral canal and develops rostro-caudally^[2]
hearing - otic capsule now dense mesenchyme. Otic vesicle elongated along the dorso-ventral axis and differentiated into the end lymphatic appendage and cochlear duct.^[3] Otic vesicle vestibular part wall thins prior to semicircular duct appearing. Geniculate ganglion forms. Auditory ossicles, tubotympanic recess and chorda tympani appear. First pharyngeal groove (cleft or hyomandibular groove) begins to form the concha and the external acoustic meatus. Six auricular hillocks present (1 tragus, 2 and 3 crus helicis, 4 and 5 helix, and 6 antitragus).
smell olfactory nerve fibres enter the brain^[4] The olfactory nerve is organized into two plexuses, lateral and medial, the latter mingled with the terminal-vomeronasal complex.^[5] Stages Description | primitive olfactory bulb

Vision - Retinal pigment is visible and the retinal fissure is largely closed. Eyelids grooves deepen, eyelid folds develop, first below, and then above, the eye.^[6]
eyelid sulcus (groove) above and below eye deepen and eyelid folds develop (below first and then above)^[6]
diaphragm - pleuroperitoneal fold (PPF) no longer separated from the diaphragm (CRL 14mm)^[7]
Abdominal Wall muscle cells now migrated approximately 50% of the distance to the ventral midline, inner and outer layers were not discernible yet.^[8]
cardiovascular
- coronary circulation acquires coronary sinus connection.^[9]
endocrine^[10]
- Hypophysis - juxtacerebral wall of the craniopharyngeal pouch is the thicker. The lateral lobes (future infundibular, or tuberal, part) and the anterior chamber (Vorraum) are clearly visible (O'Rahilly 1973 a). The infundibular recess displays a characteristically folded wall, namely the neurohypophysis (O'Rahilly 1973 a).
- thymus - connection of the thymus with the pharynx has been severed (Weller 1933). The thymus is intimately approximated to the cervical duct (ibid.) According to Norris (1937), both third and fourth pouches make contact with the ectoderm, although only the third "receives an increment from the ectoderm".
- parathyroid - parathyroid 4 is attached to the lateral surface of what Weller (1933) termed the "lateral thyroid component"
- thyroid The lobes of the thyroid curve around the carotid arteries and are connected by a delicate isthmus. Lacunae "should not be confused with lumina of follicles" (Weller 1933).
- adrenal Cortex - dorsal part of the whole suprarenal primordium is disorganized by the invasion of sympathetic nerves and cells, while the band of C2 cells and the coelomic epithelium remain intact (Crowder 1957).
- adrenal Medulla - first neural migration is at its height. Growth of the para-aortic complex is extensive. The plexiform complex is derived from paravertebral sympathetic ganglia T6-12 and usually L 1. Included in it are the primordia of the suprarenal medulla and of the celiac, superior mesenteric, and renal plexuses. Nerve fibres and "paraganglion" (M3) cells enter.
- pancreas - ventral pancreas has now fused with dorsal (Streeter 1948). Perhaps the ventral and dorsal ducts have begun to blend (Russu and Vaida 1959).
meninges (spinal cord) - vertebral rudiments stand out more and more distinctly from the intermediate zone. Over the dorsal surface of the spinal cord the closure membrane is separating into a peripheral and denser body-wall portion and a deeper and looser portion. The latter becomes part of the meninx primitiva, and this can be identiﬁed everywhere about the cord. The meninx primitiva has broadened, and its cells are more scattered than in the previous age group. It is deepest ventral to the neural tube and in the lateral parts between adjacent ganglia. The ganglia have migrated so that their ventral tips lie at the level of origin of the ventral nerve root. The vascular channels now surround the spinal ganglia, and small vessels penetrate into the spinal cord. A frontal section through the ganglia shows extensions of the dense lateral concentrations of the vertebral canal, passing medially between each two adjacent ganglia to become continuous with the meninx adjacent to the cord. These extensions are the ﬁrst indications of the denticulate ligaments of the pia mater. These primordia of the dentate processes are also observed in transverse sections, forming a cellular concentration between the ventralateral part of the neural tube and the ventral part of the neural arch rudiments.^[11]

limb - neural the median nerve, the radial nerve, and the ulnar nerve enter into the hand plate.^[12]

References

↑ Müller F & O'Rahilly R. (1989). The human brain at stage 17, including the appearance of the future olfactory bulb and the first amygdaloid nuclei. Anat. Embryol. , 180, 353-69. PMID: 2802187
↑ Patelska-Banaszewska M & Woźniak W. (2004). The development of the epidural space in human embryos. Folia Morphol. (Warsz) , 63, 273-9. PMID: 15478101
↑ Toyoda S, Shiraki N, Yamada S, Uwabe C, Imai H, Matsuda T, Yoneyama A, Takeda T & Takakuwa T. (2015). Morphogenesis of the inner ear at different stages of normal human development. Anat Rec (Hoboken) , 298, 2081-90. PMID: 26369281 DOI.
↑ Müller F & O'Rahilly R. (2004). Olfactory structures in staged human embryos. Cells Tissues Organs (Print) , 178, 93-116. PMID: 15604533 DOI.
↑ Bossy J. Development of olfactory and related structures in staged human embryos. (1980) Anat. Embryol., 161(2);225-36 PMID 7469043
↑ ^6.0 ^6.1 Pearson AA. The development of the eyelids. Part I. External features. (1980) J. Anat.: 130(1): 33-42. PMID 7364662 PDF
↑ Clugston RD, Zhang W & Greer JJ. (2010). Early development of the primordial mammalian diaphragm and cellular mechanisms of nitrofen-induced congenital diaphragmatic hernia. Birth Defects Res. Part A Clin. Mol. Teratol. , 88, 15-24. PMID: 19711422 DOI.
↑ Nichol PF, Corliss RF, Yamada S, Shiota K & Saijoh Y. (2012). Muscle patterning in mouse and human abdominal wall development and omphalocele specimens of humans. Anat Rec (Hoboken) , 295, 2129-40. PMID: 22976993 DOI.
↑ Hutchins GM, Kessler-Hanna A & Moore GW. (1988). Development of the coronary arteries in the embryonic human heart. Circulation , 77, 1250-7. PMID: 3286038
↑ O'Rahilly R. The timing and sequence of events in the development of the human endocrine system during the embryonic period proper. (1983) Anat. Embryol., 166: 439-451. PMID 6869855
↑ Sensenig EC. The early development of the meninges of the spinal cord in human embryos. (1951) Contrib. Embryol., Carnegie Inst. Wash. Publ. 611.
↑ Shinohara H. Naora H. Hashimoto R. Hatta T. and Tanaka O. Development of the innervation pattern in the upper limb of staged human embryos. (1990) Acta Anat (Basel) 138: 265-269. PMID 2389673