BIO 30 LAB: Pattens 72-hr Chick Summary 

 
I. External Features 
● Torsion – effected throughout entire length of the embryo 
○ 3  days:  complete  well  posterior  to  the  level  of  the  heart,  but  the  caudal  portion 
of the embryo is not yet completely turned on its side 
○ 4  days:  entire  body  has  been  turned  through  90  degrees,  with  left  side  on  the 
yolk 
● Flexion – cranial and cervical flexure which appeared in the 2​nd​ day have increased 
○ Nearly right-angled bends in the midbrain and in the cervical region 
○ Midbody  region  of  3-day chicks is slightly concave dorsally because the embryo 
is still broadly attached to the yolk in that region 
○ 4-day  chicks  are  only  attached  to  the  yolk  by  the  slender  yolk  stalk,  since  the 
body  folds  have  undercut  the  embryo → when the yolk stalk elongates, it allows 
the embryo to become straight in the middorsal region and then convex dorsally 
○ Caudal flexure becomes more pronounced 
○ Originally  straight  long  axis  becomes  C-shaped  and  its  head  and  tail  lie  close 
together  →  result  of  the  increase  in  flexure  of  cranial,  cervical,  dorsal,  and 
caudal regions 
● Branchial Arches and Clefts 
○ Fourth  branchial  cleft  has  appeared  caudal  to  the  three  already  formed  in 
55-hour chicks 
○ Thicker  and  more  conspicuous  than  in  younger  embryos;  aortic arches become 
more difficult to see 
● Oral Region 
○ Pharyngeal  region  and  ventral  surface  of  the  head  lie  closely  together  due  to 
cranial and cervical flexure 
○ Mandibular arch forms the caudal boundary of the oral depression 
○ Maxillary  processes,  paired  elevations  at  either  side  of  the  lateral  part  of  the 
mandibular  arch,  grow  mesiad  and  form  the  cephalo-lateral  boundaries  of  the 
mouth opening 
○ Nasal  pits  appear  as  shallow  depressions  in  the  ectoderm  of  the  rostral  part  of 
the head which overhangs the mouth 
○ Naso-lateral  process  and  naso-medial  process  surround  each  nasal  pit  in  a 
U-shaped elevation with limbs directed toward the oral cavity 
■ Naso-medial  processes  will  grow  toward  the  mouth  and  meet  the 
maxillary processes which are growing in from either side → fusion of the 
naso-medial  processes  and  maxillary  processes  gives  rise  to  the  upper 
jaw (maxilla) 
○ Fusion  in  the  midline  of  the  right  and  left  components  of  the  mandibular  arch 
give rise to the lower jaw (mandible) 

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 ● Allantois  –  small  and  still  concealed  by  the  posterior  appendage-buds  in  3-day  chicks; 
by  the  4​th  day,  it  has  undergone  rapid  enlargement  and  projects  from  the  umbilical 
region as a stalked vesicle 
● Appendage-Buds – both anterior and posterior appendage-buds are well-established 
○ Anterior (wing) bud – formed opposite somites 15-20 
○ Posterior (leg) bud – formed opposite somites 27-32 
○ By  the  4​th  day,  they  have  increased  in  size  considerably  to  form paddle-shaped 
extensions from the sides of the body 
○ Main  mass  of  buds  is  composed  of  closely  packed  mesenchymal  cells,  with 
outer covering of ectoderm 
○ Apical  ectodermal  ridge  –  thickened  band  of  ectodermal  covering  along  the 
convex outer margin of the buds 
○ 3  days:  earliest  indication  of  impending limb formation → heightened rate of cell 
proliferation  in  the  somatic  layer  of  the  lateral  mesoderm  where  the 
appendage-buds are supposed to appear 
○ Interactions between the mesodermal core and the ectodermal apical ridge 
■ Cells  emerge  from  the  parent layer of the region of appendage-buds and 
become mesenchymal 
■ Mesenchymal  cells  migrate  laterally  and  constitute  the  core  of  the 
appendage-bud 
■ Mesenchymal  cells  induce  the  formation  of  the  apical  ridge  in  the 
overlying ectoderm 
■ Ectodermal  apical  ridge,  once  established,  exercises  a  return  controlling 
action on the mesenchymal core of the bud 
∘ If  the  apical  ridge  is  removed,  the  distal  part  of  the  appendicular 
skeleton  fails  to  develop,  such  as  wing  bones  for  anterior  or  foot 
bones for posterior 
∘ The  earlier  in  development  the  apical  ridge  is  removed,  the 
greater the deficiency in development 
∘ Appendicular girdles are not affected, only distal portions 
○ Molding is also brought about by selective cell death 
■ Cell deterioration plays a huge role in the shaping of digits or wing parts 
 
II. Nervous System 
Summary of Development Prior to the Third Day 
● 16-18 hrs​: earliest indication of CNS formation 
○ Thickening of ectoderm forms neural plate  
○ Neural plate becomes longitudinally folded to form neural groove  
○ Fusion of margins of neural folds (cephalic then caudal)  
■ Neural groove closes to form tube and separates from body ectoderm  
■ Cephalic portion of tube becomes dilated to form brain  
■ Remainder: spinal cord 
● Early stages of brain: series of enlargements in ventral and lateral walls  

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 ○ Fundamental metameric structure  
○ Establishment of 3 vesicle condition 
■ Lines  of  demarcation  between  prosencephalon,  mesencephalon,  and 
rhombencephalon  formed  by  exaggeration  and  obliteration  of  some 
inter-neuromeric constrictions  
○ Original constrictions persist longest in rhombencephalon  
● 3 vesicle condition is transitory 
○ 40 hrs​: division of rhombencephalon is clearly indicated  
○ Division  of  prosencephalon  &  establishment  of  5  vesicle  characteristic  takes 
place later  
● 55  hrs​:  cranial  flexure  →  bending  of  brain;  entire  prosencephalon  is  displaced  ventrad 
and caudad toward heart  
○ Head of embryo has undergone torsion, lies with left side on yolk  
○ Flexion  and  torsion  have  changed  relationships  of  the  brain, but regions are still 
evident  
○ Prosencephalon: noticeably enlarged cephalic to optic vesicles  
■ Slight constriction in dorsal wall  
■ Demarcation begins 
 
Formation of Telencephalic Vesicles 
● End  of  3rd  day  (48 hrs)​: antero-lateral walls of primary forebrain → evaginated to form a 
pair of vesicles (each lies on either side of the midline)  
○ Lateral evaginations: telencephalic vesicles 
● Foramino of Monro 
○ Opening  through  which  cavities  are  continuous  with  lumen  of  median  brain 
portion 
● Telencephalic division of the brain includes:  
○ 2 lateral vesicles 
○ Median portion of brain from which they arise  
● Lumen of telencephalon has 3 divisions:  
○ Median telocoel (confluent posteriorly with…)  
○ Diocoele 
○ 2 lateral vesicles, connecting with telocoel via foramina 
● Median  anterior  wall  of  telocoel  (previously  most  rostral,  remains  most  rostral  part  of 
brain lying in midline) → lamina terminalis  
● Telencephalic  vesicles  →  cerebral  hemispheres;  cavities  →  become  paired  lateral 
ventricles of adult brain 
○ Hemispheres  undergo  enlargement  later  and  extend  dorsally  and posteriorly (as 
well as rostrally)  
○ Eventually  covers  entire  diencephalon  and  mesencephalon  under  posterior 
lobes  
○ Contain brain-centers for memory and actions conditioned by past experience  
● Arbitrary lines of division between adjacent brain regions  

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 ○ Telencephalon  &  diencephalon:  line  from  ​velum  transversum  to  the  ​recessus 
opticus  
■ Velum  transversum​:  ​55  hrs​;  internal  ridge  formed  by deepening of dorsal 
constriction; indicates impending division of primary forebrain  
■ Recessus  opticus​:  transverse  furrow  in  the  floor  of  the  brain;  leads  on 
either side into the lumina of the optic stalks  
● Located  rostral  to  the  optic  chiasma  where  some  optic  nerve 
fibers cross  
● Preoptic recess  
Diencephalon 
● Lateral  walls  -  little  differentiation  except  ventrally where the optic stalks merge into the 
walls of the brain 
● Epiphysis  -  median  evagination  in  the  roof;  does  not  differ  from  55  hr  except  for  some 
enlargement 
● Infundibulum 
○ depression in floor; close proximity to Rathke’s pocket 
○ Fuses with Rathke’s pocket to form ​hypophysis  
● Later, lateral walls become thickened to form ​thalamus 
○ Reducing  the  size  and  changing  the  shape  of  diocoel  (adult  anatomy:  3rd  brain 
vesicle) 
● Anterior Choroid Plexus 
○ anterior vascular vessels that invaginate the roof and push into the third ventricle 
● Tuberculum Posterius 
○ Rounded elevation in the floor of the brain 
○ Marking boundary between diencephalon and mesencephalon 
Mesencephalon 
● No specialization beyond thickening of its walls 
● Dorsal walls rapidly thickens → ​Corpora Quadrigemina 
○ Four symmetrically placed elevations 
○ Anterior pair (superior colliculi) – brain center for visual reflexes 
○ Posterior pair (inferior colliculi) – center for auditory reflexes 
● Floor of mesencephalon also thickens → ​Crura Cerebri 
○ Main pathway of fiber tracts 
○ Connect cerebral hemispheres with posterior part of brain and SC 
● Mesocoel  reduced  by  thickening  of  walls  into  a  narrow  canal,  ​cerebral 
aqueduct/aqueduct of Sylvius 
Metencephalon 
● Boundary  between  mesencephalon  and  metencephalon  -  original  inter-meuromeric 
constriction  
● Caudal boundary of metencephalon not defined 
○ Located approximately where the brain roof changes from thick to thin 
● Later, ventral and lateral extensive ingrowth of fiber tracts 
○ Gives rise to ​pons and cerebellar peduncles 

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 ● Roof  undergoes  enlargement  and  becomes  ​cerebellum  (coordinating  center  for 
complex muscular movements 
Myelencephalon 
● Dorsal wall reduced in thickness → ​thin roof of medulla  
○ Carries  the  roof  with  them  and  grows  into  the  myocoel  to  form  the  ​posterior 
choroid plexus 
● Ventral and lateral walls → ​floor and side walls of medulla  
○ Medulla:  conduction  path  between  cord  and  brain;  reflex  center  for  involuntary 
activities (breathing) 
Ganglia of the Cranial Nerves 
● Cells derived from cephalic portion of neural crest → aggregate to form ganglia 
● 96  hours​:  largest  and  most  clearly defined ganglia is the ​semilunar (Gasserian) ganglion 
of the fifth (trigeminal) cranial nerve 
● Immediately  cephalic  to  the  auditory  vesicle:  mass  of  neural  crest  cells;  primordium  of 
the ganglia of the 7th and 8th nerves  
● Posterior to auditory vesicle: superior ganglion of the 9th nerve  
● Ganglion 10/Vagus nerve 
○ Recognized in sections of chicks at end of 4th day 
Spinal Cord 
● Exhibits an elliptical lumen in cross section 
● Progression: lateral walls greatly thicken in contrast with dorsal and ventral walls 
● Lumen becomes compressed laterally until it appears as a vertical slit   
 
Thin dorsal wall  Roof plate 

Thin ventral wall  Floor plate 

Thickened side walls  Lateral plate 

 
Spinal Nerves 
● Adult – each spinal nerve connected with the cord by two roots  
○ Dorsal root – pathway for sensory (afferent) nerve fibers  
○ Ventral root – pathway for motor (efferent) nerve fibers 
● Lateral to the cord, the 2 roots unite 
● Spinal  ganglion  (dorsal  root  ganglion)  –  located  on  the  dorsal root between DC and the 
point where the 2 unite 
● Distal to the union of dorsal and ventral roots is a branch: ​ramus communicans  
○ Extends ventrad to a ganglion of the sympathetic nerve cord  
● When first formed from NCC, the spinal ganglion has no connection with the cord  
● Dorsal  root  -  established  by  growth  of  nerve  fibers  from  cells  of  the  spinal  ganglion 
mesiad into the dorsal part of the lateral plate of the cord  
○ At  the  same  time,  fibers  grow  distad  from these cells to form the peripheral part 
of the nerve  

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 ○ Fibers that arise from the DRG conduct sensory impulses toward the cord  
● Ventral  root  -  formed  by  fibers  which  grow  out  from  cells  located  in  the  ventral  part  of 
the lateral plate of the cord 
● Sympathetic  ganglia  arise  from  NCC,  and  partly  from  cells  moving  out  from  the  spinal 
cord 
○ Developing  sympathetic  ganglia:  local  enlargements  on  paired  cordlike 
structures, the ​prevertebral sympathetic chains 
○ Each  sympathetic  ganglion  is  connected with spinal nerve by ​fibro-cellular cord​, 
the primordium of the ramus communicans  
○ From  certain  points  of  the  sympathetic  ganglia, cells migrate still further ventrad 
→ establish primordia of ​splanchnic sympathetic system 
 
III. The Sense Organs 
The Eye 
● Primary optic vesicles: ​30 hrs​; dilations in the lateral walls of the prosencephalon 
● Optic  vesicles  first  open  broadly  into  the  brain,  but  later  develop  constrictions  which 
narrow their attachment to the form of a stalk 
● 55  hrs​:  primary  optic  vesicles  invaginate  to  form  double-walled  secondary  optic 
vesicles/optic cups  
○ Ventral wall is incomplete, gap called ​choroid fissure 
● Lens:  thickening  of  the  superficial  ectoderm,  becomes  depressed  to  form  a  vesicular 
invagination extending into optic cup   
● We can identify the beginning of the structures of the adult eye  
○ Thickened  internal  layer  of  optic  cup  sensory  layer  of  retina  ▪  Fibers arise from 
nerve cells in this layer of the retina  
■ Grow  along the groove in the ventral surface of the optic stalk toward the 
brain  
■ Form the optic nerve  
○ External layer → pigment layer of the retina  
○ Mesenchymal  cells  aggregate  in  progressively  increasing  number  outside  the 
optic cup 
■ Sclera and choroid coat are derived 
■ Some  make  their  way  into  the  optic  cup  thru  choroid  fissure  →  cellular 
elements of vitreous body 
○ Margins of optic cup adjacent to lens → ​complex ciliary apparatus  
○ Superficial ectoderm overlying the eye → ​corneal and conjunctival epithelium   
○ Mesenchymal  cells between lens and corneal epithelium → ​substantia propria of 
the cornea 
The Ear 
● 1st to appear: auditory placode  
○ 36 hrs​: visible as thickened plate of ectoderm 
○ Sinks below the level of surrounding ectoderm; forms the ​floor of the auditory pit 

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 ● Endolymphatic  duct:  tubular  stalk  adherent  to  superficial  ectoderm  for  a  time,  marks 
location of original invagination 
The Olfactory Organs 
● 3rd and 4th day: pair of depressions in head ectoderm 
● Located ventral to telencephalic vesicles and anterior to mouth  
● Epithelium  lining  the  pits  eventually  come  to  lie  at  the  extreme  upper  part  of  the  nasal 
chambers and constitutes the olfactory epithelium 
● Nerve  fibers  grow  from  these  cells  to  telencephalic  lobes  of  the  brain  →  olfactory 
nerves 
 
IV. The Digestive and Respiratory Systems 
Summary of Development Prior to the Third Day 
● Primary endoderm that gives rise to epithelial lining of digestive and respiratory systems 
and associated glands are established 
○ Attached peripherally to yolk 
○ Primitive gut cavity - bounded dorsally by endoderm; ventrally, yolk is temporary 
floor 
○ Only that within the embryonal area forms the ​enteric tract 
○ Peripheral: formation of yolk-sac 
● Foregut​: first part of gut to acquire complete endodermic lining  
○ Floor formed by caudally progressing concrescence of endoderm 
● Hindgut: later formed in a similar manner 
○ Progress of subcaudal fold toward head 
■ Between foregut and hindgut, the midgut remains open to yolk ventrally 
■ As embryo is more completely separated from yolk, the foregut and 
hindgut increase in extent, at the expense of the midgut   
Establishing of the Oral Opening 
● Younger chicks: when gut is first established, it ends as a blind pocket both cephalically 
and caudally 
● 55 to 60 hrs​: processes leading toward establishment of oral opening clearly indicated  
● Midventral evagination of pharynx – established immediately cephalic to the mandibular 
arch  
○ Opposite this and growing in to meet it, the stomodeal depression has been 
formed o  
○ Bar to opening: oral plate  
■ Thin membrane  
■ Pharyngeal endoderm + stomodeal ectoderm  
● 72 hours​: communication foregut with outside is establish via breaking of oral plate 
○ Rupture of oral plate growth of surrounding structures deepens originally 
shallow stomodeal depression 
○ Region where oral plate originally was: becomes the region of transition from 
oral cavity to pharynx 
● Formation of the oral opening does not take place at extreme anterior end of foregut 

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 ● Small gut pocket extends cephalic to the mouth (pre-oral gut) 
○ Rapidly becomes less conspicuous after rupture of OP 
○ Later marked by small depression (Seessel’s pocket) which disappears too 
○ Importance: margin indicates for a time where ectoderm and endoderm became 
continuous in forming the oral opening 
The Pharynx 
● Caudal to oral opening of – foregut becomes flattened dorsoventrally and widened 
laterally  
● Either side of pharyngeal lumen: extensions/bays pharyngeal pouches  
○ Each lies opposite an external gill furrow/branchial groove  
○ Leaves only a thin layer of tissue separating the pharyngeal lumen from the 
outside, composed of: 
■ Internal pharyngeal endoderm  
■ External ectoderm of bottom of branchial groove  
■ There may be mesenchyme between these 2 epithelial layers of the gill 
plate  
■ Sometimes cephalic gill plates break through → transitory open gill 
clefts 
○ Branchial arches: ​lateral walls are thickened and filled with mesenchymal cells 
between adjacent gill grooves/clefts 
■ Aortic arches lie embedded  
The Pharyngeal Derivatives 
● Several structures that do not become part of the digestive system arise in the 
pharyngeal region  
● Origin of epithelial portions from foregut endoderm and early association with this part 
of the gut tract make it convenient to connect them to the digestive system  
● Thyroid gland – arises from floor of pharynx  
○ Median diverticulum  
○ Appears at cephalo-caudal level between 1st and 2nd pair of pharyngeal 
pouches  
○ Later: moves caudad in the body, but retains connection with root of the tongue 
via narrowed ​thyroglossal duct​ for a time 
● Postbranchial bodies - paired evaginations, arise posteriorly from 4th pharyngeal 
pouches 
● Parathyroid glands – two pairs; arise as bud-like outgrowths from the pharynx  
○ One pair is budded off the caudal faces of the 3rd pharyngeal pouches  
○ Other pair: fourth pouches  
○ Parathyroid primordia not recognizable until later 
● Thymus – barely indicated on 4th day  
○ Originates from diverticula arising from posterior faces of 3rd and 4th 
pharyngeal pouches  
○ Original epithelial character → lost in extensive ingrowth of mesenchyme  

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 ○ Organ becomes lymphoid in histologic characteristics  
The Trachea 
● 72 hrs​: 1st indication of the formation of the respiratory system  
○ Midventral groove in pharynx (​laryngotracheal groove​)  
○ Beginning posterior to the level of the 4th pharyngeal pouches and extends 
caudad  
○ Deepens rapidly and becomes separated from pharynx via closure of dorsal 
margins (except at cephalic end)  
○ Tube formed: ​trachea 
○ Opening that persists between laryngeal end of trachea and pharynx: ​glottis  
○ Original endodermal evagination gives rise only to the epithelial lining of the 
trachea 
○ Supporting structures of the tracheal walls derived from the surrounding 
mesenchyme   
The Lung-buds 
● Tracheal evagination grows caudad and bifurcates to form a pair of ​lung-buds 
● As they develop, they grow into loose mesenchyme on either side of midline 
● Relationship of mesenchyme to growing endodermal epithelium (also seen in digestive) 
is interesting  
○ Formation of tubular structures (bronchi or ducts and secreting tubules of 
digestive): dependent on presence of mesoderm about growing endoderm  
○ Endoderm grown in culture without mesoderm: spreads out as a flat, 
unorganized sheet  
■ If mesenchymal cells are added, endodermal epithelium forms tubular 
structures  
■ Tubular structures → characteristics of lung/digestive gland/epithelial 
lining of gut tract according to primordial region where the original 
endodermal cells were taken  
● Caudo-lateral growth of primordial lung-buds: surrounding splanchnic mesoderm 
pushed ahead of them and constitutes the outer investment 
● Endodermal buds give rise only to epithelial lining of bronchi and air passages & air 
chambers of the lungs  
● Connective tissue stroma of lungs: mesenchyme immediately surrounding the 
endodermal buds  
● Pleural covering: primary investment of splanchnic mesoderm  
The Esophagus and Stomach 
● Caudal to glottis: narrowed region of foregut → esophagus  
● Farther caudally: slightly dilated region → stomach  
● Concentration of mesenchymal cells about the endoderm of the esophageal and gastric 
regions → muscular and connective tissue coats 
The Liver 

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 ● Arises as diverticulum from the ventral wall of the gut, a little caudal to the stomach  
● 22 somite stage​: ​hepatic primordium​ first recognizable  
○ Appears as part of the gut from which it arises acquires a floor by the 
concrescence of the margins of the anterior intestinal portal  
○ Evagination (​primordium of the liver​)  
■ Located shortly on the lip of the intestinal portal 
■ Grows cephalad towards the confluence of the omphalomesenteric veins 
before they enter the sinus venosus 
○ Complete closure of gut floor: ​hepatic diverticulum​ lies in characteristic position 
at ventral wall of gut  
● Liver, from its first appearance, is invested by mesoderm 
● Proximal portion of original evagination: open to intestine and serves as duct of the liver  
○ Primitive duct later differentiates  
○ Adult: common bile duct, hepatic and cystic ducts, and gallbladder.  
○ Cellular cords which bud off from diverticulum → secretory units of liver (hepatic 
tubules) 
● Same process of concrescence that closes the floor of the foregut involves the proximal 
portion of the omphalomesenteric veins 
● As AIP moves caudad in lengthening of foregut, the proximal portions of the 
omphalomesenteric veins are brought together in the midline and fuse  
○ Fusion extends caudad nearly to the level of the yolk-stalk  
○ Distal to this point, they retain original paired condition  
● Liver surrounds the fused portion of the OV in its growth  
○ This early association foreshadows how blood channels coming from the 
vitelline vascular plexus become involved in the establishment of the 
hepaticoportal circulation of the adult  
The Pancreas 
● Derived from evaginations appearing in the intestinal walls in the same level as liver 
diverticulum  
● 3 pancreatic buds:  
○ Single median dorsal  
■ 72 hrs  
■ Arises from gut endoderm directly opposite the liver diverticulum and 
grows into dorsal mesentery  
○ Pair of ventrolateral  
■ Towards end of 4th day  
■ Arise close to point where duct of the liver connects with the intestine  
■ Ducts of liver and central pancreatic ducts open into the intestine by a 
common duct (​ductus choledochus​)  
○ Later, the masses of cellular cords derived from these grow together and 
become fused into a single glandular mass  
○ In birds, usually 2 or sometimes all 3 persist 
The Midgut Region 

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 ● Gut tract between stomach and yolk-stalk + anterior fourth of the gut that lies caudal to 
yolk-stalk → ​small intestine  
● Posterior two-thirds of hindgut →​ large intestine and cloaca 
The Cloaca 
● 4 days: beginning of formation indicated by dilation of posterior portion of hindgut  
● Extensive differentiation does not appear until later 
● Adult: common chamber into which intestinal contents, urine, and gametes are received 
for discharge   
● First appearance in embryo establishes the relationship of the cloaca and intestine  
● Proximal portion of allantoic stalk (​homolog of urinary bladder of mammals​): opens 
directly into cloaca 
● When urinary system is considered, the ducts that drain the developing excretory 
organs also open into the cloacal region on either side of the allantoic stalk 
● Ventral outpocketing of hindgut arises caudal to the point at which allantoic stalk opens 
into the cloaca  
● Depression appears in overlying ectoderm  
○ External depression that grows in toward the gut pocket: ​proctodeum  
● Double epithelial layer (gut endoderm + proctodeal ectoderm) → ​cloacal membrane  
● Formation of the 2 indicates the location of the future cloacal opening  
● Cloacal opening does not form at extreme posterior end of the hindgut  
● There is a post-anal pocket of the hindgut  
 
V. Circulatory System 
Interpretation of the Embryonic Circulation 
● Aortic arches 
○ Recapitulation of ancestral conditions 
○ Fundamental functional necessity of a channel connecting the ventrally located 
heat with the dorsally located heart 
○ Relationship of aortic arches to gill arches - individual development an 
unequivocal record of evolutionary phase when gills were a center of primary 
metabolic importance 
● Tendency to repeat phyletic history → earliest form of circulatory mechanism or bird or 
animal is patterned after a more primitive type; cannot be miniature of adult conditions 
● Only relatively late in development can we expect to see the emergence of the vascular 
plan characteristic of the adult 
● All changes in the circulatory system of a living organism must be gradual 
Main Routes of the Embryonic Circulation 
● Three main arcs: 
○ Vitelline arc carries blood to yolk-sac where food materials are absorbed, returns 
the food-laden blood to heart for distribution within embryo 

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 ○ Allantoic circulation is where blood gives off its carbon dioxide and acquires 
fresh oxygen supply; eliminates nitrogenous waste material from blood 
○ Intra-embryonic circulation brings food material and oxygen to, and carry waste 
material from, the various parts of developing body 
● Heart: blood of vitelline, intra-embryonic and allantoic circulations mingle 
Vitelline Circulation 
● Earliest indication of blood and blood vessel formation is at the chick’s source of food 
supply 
● Blood islands 
○ appear in extra-embryonic splanchnopleure of yolk-sac toward end of 24 hours 
of incubation 
○ Form vascular endothelium enclosing central clusters of primitive blood 
corpuscles 
● Extension and anastomosing of neighboring islands → plexus of blood channels formed 
in yolk sac 
● Extension of vitelline plexus → communication with omphalomesenteric veins 
○ Developed in embryo as caudal extensions of endocardial primordia 
● 48 hours: ​omphalomesenteric arteries establish communication between dorsal aorta 
and vitelline plexus 
○ 40th hour: ​blood cells formed in yolk-sac carried into body of embryo 
● Acquisition of food material by blood depends on activities of the endodermal cells 
lining the yolk-sac 
● Cells secrete digestive enzymes which break down deutoplasmic granules 
● Liquefied material absorbed by yolk-sac cells and transferred to blood 
● Blood carries food material in soluble form to embryo where it is assimilated 
Allantoic Circulation 
● Allantoic arteries arise by the prolongation and enlargement of a pair of vessels arising 
ventrally from the aorta at the level of allantoic stalk 
○ Blood is distributed in a rich plexus of vessels which ramify in mesoderm of 
allantois 
● Size increases rapidly as allantois increases in extent 
● Situation of allantois directly beneath porous shell: blood can carry on interchange of 
gases with outside air 
○ Plexus of small allantoic vessels where surface exposure is great 
○ Blood gives off its carbon dioxide and takes up oxygen 
● Later stage of development: ducts of embryonic excretory organs open into allantoic 
stalk near its cloacal end 
● As excretory organs become final, allantoic vesicle becomes repository for the 
nitrogenous waste materials eliminated through them 
○ Watery part of waste materials evaporates 
○ Remaining solids deposited in vesicle accumulate in extraembryonic portion of 
allantois 
○ Remain there until portion of allantois is discarded at the close of embryonic life 

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 ○ Blood from allantois collected and returned to heart over allantoic veins 
○ Distal portion of allantois: smaller veins converge and unite into 2 main vessels: 
right and left, which enter the body of embryo with allantoic stalk 
○ After entrance, allantoic veins extend cephalad in lateral body walls 
○ Enter sinus venosus on either side of OV entrance 
Intraembryonic Circulation 
● Earliest vessels to appear are large vessels communicating with heart 
● 33 hrs; ​ventral aorta leads off from heart cephalically, bifurcates ventral to pharynx give 
rise to a single pair of AA 
● AA pass dorsad around the antero-lateral walls of pharynx and continue caudally along 
dorsal wall of gut as paired dorsal aortae 
● Towards 48 hours, visceral clefts and arches appear; original pair of AA comes to lie in 
mandibular arch 
● In each visceral arch posterior to mandibular, new AA are formed connecting the ventral 
aorta with dorsal aorta 
● 50 hrs,​ 2 pairs of AA are present and 3rd beginning to form 
● 60 hrs,​ 3 AA completed, 4th starting to form 
● End of 4th day, rudimentary 5th and well-developed 6th 
● 1st and 2nd: marked diminution in size; indicative of final appearance 
● Late in 2nd day: appearance of plexiform channels extending from 1st aortic arch 
toward forebrain 
○ Foreshadow formation of internal carotid arteries 
● 60 hrs:​ ​internal carotid artery established​ as trunk extending toward brain from where 
1st AA turns into dorsal aortic root 
● 96 hrs: regression of first 2 AA 
○ Prominent role in formation of external carotid arteries 
○ Dorsal aortic root - feeder to original internal carotid artery 
■ Making it appear to originate where 3rd aortic arches merge with dorsal 
aortic root 
● Reptiles, birds, mammals: main adult vessels which connect heart with dorsal aorta are 
derived from 4th pair of AA of embryo 
○ Birds and mammals: 1 of the arches degenerates 
○ Birds: left arch degenerates; right becomes arch of adult aorta 
○ Mammals: right arch degenerates; left becomes arch of adult aorta 
● Dorsal aorta: paired at first, later become fused to form median vessel 
○ Fusion begins at cardiac level 
○ Extends cephalad shortly, never involving the dorsal aortic roots where they 
receive aortic arches 
○ Caudally, the aorta becomes fused through entire length 
● Level of anterior appendage-buds: several pairs of dorsal segmental arteries extend 
into wing buds 
○ Later, one becomes enlarged to form subclavian arteries 
● Coincident with allantois development, allantoic arteries become enlarged and extend 
into it 

[AREVALO, ROBLES]
 ○ A pair of ventral segmental vessels opposite the allantoic stalk  
○ Feed a plexus of small vessels which (with growth of allantois) come to lie 
beneath porous shell and become concerned with gaseous interchanges 
● Blood supply to posterior appendage: starts as plexus fed by several segmental vessels 
at level of appendage-bud 
○ Primordial plexus later comes to be fed by a branch arising from allantoic artery 
○ New vessel: external iliac artery 
● Adult: 3 vessels arising from dorsal aorta supply abdominal viscera 
○ Coeliac artery 
○ Superior mesenteric artery 
○ Inferior mesenteric artery 
● As development progresses, any arterial trunk growing into an organ will exhibit a 
pattern of branching which bears definite relation to the structure of the organ it is 
supplying 
● Anterior and posterior cardinal veins 
○ Principal afferent systemic vessels of the early embryo 
○ Appear toward end of the 2nd day as paired vessels extending cephalically and 
caudally on either side of midline 
○ Level of the heart: those on same side of the body become confluent in common 
cardinal veins and turn ventrad to enter SV 
● Posterior cardinals lie at first in the angle between the somites and the lateral 
mesoderm  
○ Mesonephros develops from intermediate mesoderm → PCV lie dorsal to them 
throughout length  
○ Situated ventrally in mesonephros: small irregular vessels roughly paralleling the 
PCV and anastomosing freely with them → subcardinal veins  
■ A relic of the renal portal circulation of more primitive forms 
■ Important to birds/mammals because they form the posterior vena cava 
● Young embryos: posterior cardinals are main afferent vessels of posterior part of body  
○ Later replaced by posterior vena cava 
Heart 
● Heart in adult vertebrates: ventral, unpaired structure 
● Origin in chick: paired primordia 
● After fusion of paired primordia, heart is nearly a straight double-walled tube 
○ Primordial endocardium has same structure and arises in same manner as 
endothelial walls of primitive embryonic blood vessels with which it is 
continuous 
○ Epimyocardial layer: outer investment which surrounds and reinforces 
endocardial wall 
○ Dev’t progresses: epimyocardium becomes thickened and differentiates into 2 
layers 
■ Heavy muscular layer: myocardium 
■ Thin non-muscular covering: epicardium 

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 ● Apposition of paired primordia of heart to each other: splanchnic mesoderm from either 
side of body comes together dorsal and ventral to heart 
○ Double layered supporting membranes formed 
■ Dorsal mesocardium - suspends heart in cavity; later disappears except 
at caudal end 
■ Ventral mesocardium - disappears shortly after formation 
○ Heart comes to lie in pericardial cavity unattached except at its 2 ends 
■ Cephalic end: fixed with reference to body of embryo where ventral aorta 
lies embedded ventral to floor of pharynx 
■ Caudal end: fixed by persistent portion of dorsal mesocardium and 
omphalomesenteric veins 
● 30 hrs:​ Bending of heart to form U-shaped tube begins to be apparent  
● 40 hrs​: ventricular region lies to the right  
● Bending of heart to the side involves ​mechanical expediency 
○ Initiation of bending process depends on more rapid elongation of heart than the 
chamber in which it lies fixed by its 2 ends 
○ Bending takes place to the side rather than dorsally or ventrally: attributed to 
impediment offered to its dorsal bending by the body of the embryo and ventral 
bending by yolk 
● 40 hrs​: greatest extent of lateral bending 
● Closed part of U-shaped bend becomes twisted on itself to form a loop 
○ Atrial region forced slightly to the left 
○ Truncus arteriosus thrown across atrial region by being bent to the right then 
dorsocaudad 
○ Ventricular region constituting the closed end of the loop swings dorsalward and 
toward the tail  
■ Crowded in this direction by increasing flexion of the cephalic part 
○ Original cephalo-caudal relations of atrial and ventricular are reversed  
■ Atrium now lies cephalic to ventricle  
● Atrial and ventricular regions  
○ Formerly continuous, become marked by a constriction  
○ As they become enlarged, the demarcation is accentuated  
○ Atrio-ventricular canal  
● Changes in external shape of heart → whole heart comes to occupy a more caudal 
position   
○ Heart first lies at the level of the rhombencephalon 
○ End of 4th day: tip of ventricle lies at the level of the anterior appendage-buds 
● 3rd and 4th days: changes become apparent in ventricular portions of cardiac wall 
○ Primordial tubular heart​ was first established: endothelially lined lumen was 
smooth, regular in diameter  
○ Outside the endothelium, between it and developing muscle, was a layer of 
cardiac jelly  
○ 50 – 55 hrs​: myocardium shows irregular projections extending into cardiac jelly 
→ start of ​trabeculae carneae  

[AREVALO, ROBLES]
 ○ As they grow, the endothelial lining tends to extend between them and follow 
the configuration of the muscular strands separated from them only by a thin 
layer of cardiac jelly  
○ Once established, trabeculation shows richly branching pattern  
○ Ventricular wall becomes honeycombed by tortuous intertrabecular spaces  
■ All endothelially lined and communicate (directly/indirectly) with main 
lumen of the ventricle  
■ Brings blood in close relationship to the growing cardiac muscle during 
the period before the coronary circulation to the myocardium has been 
formed  
○ Human embryos: heart is beating and effectively propelling blood through 
embryonic vessels for about a month before coronary vessels develop  
■ During this period, myocardium is entirely dependent on blood that 
reaches it by the intertrabecular spaces  
● 4th day: primordial epicardium consists of single layer of cuboidal cells  
○ Forerunner of mesothelium of epicardium  
■ Later, a supporting layer of epicardial connective tissue will form 
between the mesothelium and underlying myocardium 
● Loosely aggregated masses of mesenchymal cells lying between the endocardium and 
myocardium in the region of atrioventriular canal and of ventricle opening into truncus 
arteriosus 
○ Evidence that some of these cells arise from endothelium instead of all being 
from myocardium 
○ They all move into the space between 2 primordial layers of heart using ​cardiac 
jelly ​as a substratum for migration 
○ When aggregated, they constitute the ​endocardial cushion tissue​,  
■ Later takes part in partitioning the atrio-ventricular canal and forming the 
connective tissue framework of cardiac valves 
● Physiologic differences in myocardium of basic cardiac regions  
○ Establishment of cardiac tube - sequential manner of formation  
○ As foregut acquires a floor → cardiac primordia meet and fuse in the midline  
○ Truncoventricular part of heart is formed first → fusion reaches atrial region → 
SV is established caudal to atrium  
○ Pulsation of myocardium appears in these regions in the same sequence they 
are laid down  
○ 1st contractions: ventricle, before atrium is fully established; slow 
○ When atrium begins to pulsate, heart rate increases 
■ Transection experiments demonstrate faster rate of pulsation of atrial 
myocardium, takes control of cardiac rhythm as pacemaker  
○ After formation of SV caudal to atrium: rate of contraction > atrium  
○ Cephalo-caudal gradient in the intrinsic rate of myocardial contraction  
○ Pacemaking center = intake end   
 
VI. Urinary System 

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General Relationship of Pronephros, Mesonephros, Metanephros 
● Pronephros 
o Most anterior 
o First to be formed 
o Wholly vestigial 
o Most primitive of the vertebrate stock 
● Mesonephros 
o Principal organ of excretion during early embryonic life but disappears in the 
adult except for parts of its duct system associated with reproductive organs 
o Homologous with adult excretory organs of fishes and amphibia 
o Formed later than pronephros and formed caudal to it 
● Metanephros 
o Most caudally located of the excretory organs 
o Last to appear 
o Functional toward end of embryonic life when mesonephros is disappearing 
o Persists permanently as the functional kidney of the adult 
● All three: 
o Derived from intermediate mesoderm 
o Tubular in nature 
o Collect waste materials from capillary plexuses  
● Pronephros empties into pronephric duct 
● Pronephric ducts: 
o Formed at level of pronephric tubules  
o Extend caudad and open into cloaca 
o Close proximity with developing mesonephric tubules 
● Mesonephric tubules extend toward pronephric ducts and soon open into them 
● Pronephric tubules begin to degenerate 
● Ducts that originally arose in connection with pronephros are appropriated by 
developing mesonephros  
o Degeneration of pronephric tubules → same ducts are called mesonephric 
ducts  
Pronephric Tubules of the Chick 
● Pronephros  
o Represented by tubules which first appear at ​36 hours  
o Arise from intermediate mesoderm, or nephrotome, lateral to somites 
▪ Paired, segmentally arranged structures 
▪ A tubule appearing on either side opposite each somite from the 5​th​ to 
16​th  
o 38 Hours: ​Section passing through 10​th​ to 14​th​ somites show pronephric tubules 
favorably 
o Each tubule arises as a solid bud of cells from intermediate mesoderm near its 
junction with lateral mesoderm 

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Mesonephric Tubules 
● Develop from intermediate mesoderm caudal to pronephros 
● Early steps in formation are well shown in transverse sections of 29 to 30 somites or ​55 
hours 
● Posterior region is less advanced than they are more anteriorly 
● Appear first as cell clusters formed in intermediate mesoderm ventro-mesial to cord of 
cells or primordium of pronephric duct 
● 55 hours​: primordial cell cord representing pronephric duct → hollowed out to establish 
a definite lumen 
● Most anterior of mesonephric tubules also acquired a lumen 
● More posterior mesonephric tubules do not become associated with duct until later, but 
remain as a series of isolated vesicles 
● Mesonephric tubule differs from pronephric tubules chiefly in its relation to the blood 
vessels associated with it 
o Develops a cup-like outgrowth into which a knot of capillaries is pushed 
▪ Glomerular capsule/Capsule of Bowman: cup-shaped outgrowth from 
tubule 
▪ Glomerulus: tuft of capillaries 
▪ Glomus: small, localized capillary tufts from the continuous vascular 
ridge; diminutive form glomerulus 
● Regulated resorption of substances in nephric tubules is responsible for maintenance of 
delicate salt-water balance which exists in blood and tissue fluids 
● In the process, waste materials are left within tubule and carried by balance of water 
into excretory duct and into allantois for storage 
● Mesonephros is made up of tubules which form no nephrostome but depend entirely on 
their glomerular apparatus for their fluid intake 
 
VII. The Coelom and Mesenteries 
● Body cavity  
o Formed by the partitioning off of the primary body cavity or embryonic coelom 
o Three regions: pericardial, pleural peritoneal 
▪ Pleural cavities – paired; laterally situated sac containing one of the lungs 
▪ Pericardial chamber - contains heart 
▪ Periotoneal chamber - contains viscera, other than lungs and heart, are 
unpaired 
● In the chick, coelom: 
o Arises by splitting of lateral mesoderm of either side of body 
o Primarily a paired cavity 
o Does not show segmental pouches corresponding with somite arrangement  
● Left and right coelomic chambers extend cephalo-caudally through entire lateral plates 
of mesoderm 
● Coelomic chambers: 
o Extend on either side into mesoderm 

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 o Spreads out over yolk surface 
● Large part of primitive coelomic chambers become extra-embryonic 
● Coelom that become embryonic body cavities is marked off by series of folds which 
separate embryo from yolk 
● As ventral body-wall closes, embryonic coelom becomes completely separated from 
extra-embryonic 
● Delayed closure of ventral body-wall in yolk-stack region results in embryonic and 
extra-embryonic coelom retaining communication for a time after they have been 
completely separated elsewhere 
● Ventral body-wall folding process also completes gut ventrally  
● Right and left coelomic chambers are expanded mesiad 
● Newly closed gut lie suspended between double layer of splanchnic mesoderm (primary 
mesentery) 
o Constitute mesial walls of right and left coelomic chambers 
o Apposed to gut 
o Supports gut in body cavity 
● Ventral mesentery: ventral to gut; attached to ventral body-wall; early disappears 
● Dorsal mesentery: dorsal to gut; attached to dorsal body wall; persists in large part 
● Established dorsal and ventral mesenteries —> complete membranous partition (left 
and right halves of body cavity) 
● Primary dorsal mesentery persists in large part but ventral mesentery early disappears 
● Gut is embedded in dorsal body wall in most cephalic part of body cavity 
● Dorsal mesocardia – primary ventral mesentery dorsal to heart 
● Ventral mesocardia – ventral to heart  
● When ventral and dorsal mesocardium break through, the primary right and left 
coelomic chambers become confluent to form the pericardial region of body cavity 
● Liver arises as an outgrowth from gut and extends into retained part of ventral 
mesentery 
● Entire dorsal mesentery persists and forms the supporting membranes of digestive tube 
o Mesogaster – dorsal mesentery which suspends stomach 
o Mesocolon – dorsal mesentery supporting colon 
TRANSER’S NOTES 
- You can read simoune’s pattens summary in the gdrive for more notes since we deleted 
irrelevant info na pang-4th day and/or mema lang for this summary (pero ang haba pa 
rin huhu) GOOD LUCK GUYS WE GOT DIZ\ 
 
 

[AREVALO, ROBLES]