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    If you remember last time we talked about the

    development of the oral region and we started talking

    about branchial arches , we said these branchial arches

    are structures that appear in the lower part of the face andalso the neck , starting from the end of week three and

    they appear as arches , as we said they appear as arches

    so this is the head of the embryo and this is the lower jaw ,

    we can see the first branchial arches , the second

    branchial arch and third branchial arch and so on

    These are six arches , each arch is separate from the arch

    below it by a groove.

    For example these groove is called the branchial cleft ,

    from inside if you look at theses groove from inside the

    embryo also you will see grooves but these grooves are

    called branchial pouches , so the groove is called pouch if

    it is from inside , it is called cleft if it's from outside , the

    arch itself is called branchial arch and we have six arches

    , it is not necessary to see all the arches at the same time

    but what you do actually you see , for example arch

    number one and two and three maybe but when you start

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    see arch number four arch number one is disappear

    because arch number one will have added to the

    development of the lower part of the face

    arches

    arch number 4

    arch number one

    arch

    Last time we said that each arch has skeletal elements ,

    nervous elements , muscular elements and vascular

    elements and we said skeletal elements of face arch aremeckels cartilage and so on , the nervous elements are

    trigeminal nerve for first , facial nerve for second ,

    glossopharngeal for third and from four to six we have the

    vagus nerve and also if you remember , each muscle that

    is supplied by one of these nerves should also originate

    within the arch , for example if we have the muscle supply

    by trigeminal nerve like master muscle or temporalismuscle this means that these muscle develops within the

    first branchial arch because this is the nerve for the first

    branchial arch .

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    And we give an example about one of the muscle that

    develops in two arches which is the digastric muscle , the

    digastrics muscle located in the floor of the mouth , it has

    two bellies , the first belly is supplied by trigeminal , by thisreason it develops in the first arch , the posterior belly is

    supplied by facial nerve , for this reason this belly is

    developed in the second branchial arch , and also we

    discussed the details of the muscular elements and the

    vascular elements and now we will come to the

    pharyngeal pouches .

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    this is how the arches look like from inside , we make a cut

    and we are looking at the arches from insidenot from outside . for this reason these grooves here by

    this one and this one and this one are not called branchialcleft because these are located inside and called branchial

    pouches so branchial pouches important for the

    development of the tongue so the tongue is develop from

    the branchial pouches .

    The first part of the tongue starts to appear at age of

    thirty two days and it develops from different swelling

    , notice first that here we have the natural swellings

    and also we have what we called tuberculum impar

    in the middle , these three swelling they are related

    to which arch ?? arch number one , so this makes

    the anterior part of the tongue , so these swelling ,

    the two lateral swelling plus tuberculum impar which

    is the medial swelling they later on fused together ,

    they swell and fuse together making the anterior two

    third of the tongue , now the swelling from the third

    arch , also we have a big swelling It's called

    Copula/Hypobranchial eminence this develops from

    arch # 3 as you see but it over grows the second

    arch it over laps the second arch

    arch 3

    arch 2

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    And what does this lead to ?this lead to the development

    of the posterior part of the tongue or the posterior third of

    the tongue

    See here this is the tongue after development , we can

    see the first part of the tongue or the anterior two third of

    the tongue which is the body of the tongue , this develops

    from the first branchial arch and regarding the posterior

    part of the tongue it develops from third branchial arch .

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    The root of the tongue which is the very much posterior

    part of the tongue which is just next to the epiglottis this

    develop from arch number 4 , so we have also swelling

    from the fourth arch which is particularly . these twoswelling or this part as we see these are the extreme

    posterior of the tongue ( root of the tongue )

    So on the exam if I said tongue develops from ?? we have

    to say arch one , arch three and arch four , these three

    gives the body of the tongue . arch one give the anterior

    two third of the tongue and arch 3 give the posterior third

    of the tongue and arch 4 give the root of the tongue .

    Do we have any contribution from the second arch ?yes

    the second arch only contribute to the taste buds so the

    taste buds of on the tongue they are derived from the

    second arch.

    For this reason because the anterior part of the tonguefrom arch # 1 , the sensation or the sensory innervation is

    from trigeminal nerve and the posterior third of the tongue

    is innervated by glossopharngeal nerve which is the nerve

    of third arch , and the root of the tongue is from the vagus

    nerve , and the taste buds of the tongue are from the

    facial nerve because it develops from arch # 2 .

    Now what happens later on ?after fusion we still see some

    v-shape sulcus so when the anterior two third of the

    tongue fuse with the posterior third of the tongue , fusion

    leads to sulcus

    Fusion 011 %

    v-shape sulcus this is called sulcus terminalis , its the

    junction between the anterior part and posterior part of the

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    tongue or if you like its the junction between the part of

    the tongue that develops from the first branchial arch and

    the part developing from the third branchial arch.

    what is located anterior the sulcus terminalis should be

    supplied by trigeminal nerve which is the cranial nerve of

    the first arch , and what is located posterior to it is

    supplied by glossopharngeal nerve .

    the very median part of the median end of sulcus

    terminalis we have what we call foramen cecum which is

    small depression , this foramen cecum is importantbecause this is the origin for the development of thyroid

    gland .

    this actually happen by a duct that drops down from this

    foramen cecum its called thyroglossal duct , why do we

    call it thyroglossal ?? because it's now within the tongue

    so glossal , thyro from thyroid so thyroglossal, this drops

    down from foramen cecum and descend until it reach the

    neck region where it swell and develop the thyroid gland.

    this means thyroid gland develops from the area between

    the anterior two third and posterior third of the tongue , forthis reason we may see thyroid tissue within the tongue ,

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    why ?? because this tissue is eminence Of the thyroid

    gland , we call it ectopic thyroid tissue , also this ectopic

    thyroid tissue is functional.

    we have group of papillae that are usually located

    anteriarly To sulcus terminalis , so these are located within

    the anterior two third of the tongue , but the origin of these

    papillae is from the posterior third , for this reason these

    taste buds that are present on the circumvallate papillae

    these are supplied by glossopharngeal nerve although

    they are located anterior to sulcus terminalis ,, why ??

    because during the development of the tongue these

    papillae migrate from the posterior third to the anterior two

    third so they cross sulcus terminalis.

    we have taste buds on vallate papillae , these taste buds

    are response for the sour taste . innervation for this

    papillae is glossopharngeal nerve

    sulcus papillae terminalis anterior two third of the tongue

    supplied by trigeminal supplied by

    glossopharngeal

    posterior third sulcus terminalis sulcus terminalis

    For this reason they take the embryological innervationwith them . the tongue sensory the first nerve which is

    trigeminal nerve provides sensory innervations for the

    anterior two third of the tongue , sure except the region of

    the circumvallate papillae , the taste in the anterior two

    third of the tongue is from the facial nerve , the taste at the

    posterior part of the tongue is from glossopharngeal nerve

    this is also include the taste buds located at the vallatepapillae , the extreme posterior of the tongue is from

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    vagus nerve which is the cranial nerve # 10 that is

    responsible for the innervations of the 4th , 5th , 6th arches ,

    so thats why the 4th arch nerve through the branch called

    superior laryngeal nerve , its supply both sensory andtaste to the root of the tongue , if we have taste buds at

    the extreme posterior part of the tongue this is supplied by

    vagus nerve , the posterior third sensory and taste from

    third branchial arch which is glossopharngeal nerve and

    regarding the motor supply , the muscles of the tongue

    they get very special innervations from another nerve , this

    nerve is called hypoglossal nerve which is cranial nerve #12 hyogloosal nerve , the innervations of the muscles of

    the tongue the intrinsic and extrinsic muscle.

    Remember the myotoms , they are from the metotic

    somites , remember last lecture we said we have the head

    somites , we have the prootic somites will form the

    muscles of the eye and the myotom of metotic somites will

    form the muscles of the tongue , so thats why metotic

    somites carry with them the hypoglossal nerve supplied

    muscle .

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    The development of the face :

    The face grows by number of process .we have the

    maxillary process , mandibular process and frontonasal

    process , together these proceses they make the face .

    Maxillary and mandibular process are paired process ,

    one process on the right and the another on the left .

    Let us see this big process , FNP this is the frontonasal

    process , this process here and here are the maxillary

    process , and this long process here is the mandibular

    process .

    Face develop around stomodeum ,stomodeum is the

    primitive mouth .

    Can you see here this depression above the mandibular

    process ?this cavity is called stomodeum or the primitive

    mouth of the embryo .

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    The primitive mouth of the embryo is separated from the

    beginning of gastrointestinal tract by the buccopharngeal

    membrane which came from prochordal plate , and we

    said before the age of 21 day this membrane is active sothis means that the stomodeum is separated from the

    gastrointestinal tract . But at the age of 21 days this

    buccopharngeal or oropharungeal membrane is rupture

    Q: the mouth is communicated with GIT tract at the age of

    1 _ 10 days ( F )

    2_ one month ( T )

    Each branchial arch is covered outside by ectoderm and

    inside by endoderm . for example arch 2 the outside

    covering is ectoderm and the covering inside is endoderm.

    the tongue in the first brachial arch from inside and

    outside is covered with ectoderm. *** question in the exam

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    **The frontonasal process is not from the brachial arch

    only the maxillary and mandible processes from firstbrachial arch.

    The frontonasal process " the area in pink " develops pits

    and it's called nasal pits later it will become nostrils and at

    the sides of these pits we find some swellings 2 one

    medial called the medial nasal prominence and the lateral

    nasal prominence . notice how big the distance isbetween the pits at the beginning after that they start to

    migrate towards each other , around each nasal pit there

    is swelling , lateral nasal swelling and medial nasal

    swelling , the 2 medial nasal process of the 2 nostrils fuse

    together forming the intermaxillary segment which form

    the tip of the nose , the collamela of the nose (area

    between the nostrils )and the median part of the upper lip.and we call it like that because it descends down and take

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    place between the 2 maxillary processes , the lateral nasal

    process remain within the frontonasal process this gives

    the mid portion of the nose and the philtrum of upper lip

    inside it there is the area of bone that carries the maxillarypalate ( contain the maxillary incisors central and lateral

    incisors ) .

    **The upper margin of the upper lip is like M but the lower

    lip is one piece and why is that ? because the median part

    of the upper lip from the frontonassal process

    Regarding to the lateral nasal swelling it becomes the alaeof the nose ( lateral to the nostril ) , the maxillary

    processes become the maxilla and the mandibular

    processes fuse together forming the mandible .

    Nasolacrimal duct :

    Its a canal that take the tears down to the nose and its

    form in the groove between the maxillary process and thelateral nasal swelling.

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    Between the maxillary process and lateral nasal swelling

    we have a groove this groove will deepen and create a

    canal called lacrimal canal or nasolacrimal duct ( joints the

    lacrimal bone with the nose ) when u start crying when ustart releasing tears the first thing to feel is running nose

    now the extra tears that the duct can't coop with will go out

    and wet your skin . so what happens if something went

    wrong the maxillary process fails to fuse with the philtrum

    of the upper lip we will have a condition called cleft lip .

    We have to define two types of two part of the palate ,

    primary palate and secondary palate , the anterior region

    of the palate that carries the central and lateral incisors is

    called the primary palate and the remain part is called the

    secondary palate , primary and secondary palate are

    formed separately and finally they fuse together forming

    the whole palate .

    Secondary palate has two parts one on the left side andone on the other side , these are called the palatine

    processes , at first these processes are vertical , why are

    they vertically oriented not horizontal ? because we have

    a structure that occupies the space between them , this

    space is occupied by the tongue , with time , with the

    facial growth , the tongue becomes lower and drops in the

    space between the two palatine processes , which allowsthe processes to go horizontally and fuse with the primary

    palate , so now we have primary palate anteriarly and two

    palatine processes posteriorly , the three part fuse

    together and form the whole palate , thats why they start

    to adjust themselves ( they start to go horizontal instead of

    vertical )

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    In anatomy the palate is formed by two bones the maxilla

    and the palatine bones , here we talk about separate

    processes and not separate bones , that help in forming

    the palate , so , the tongue is in between , horizontalreorientation and then they start to reorient themselves ,

    the tongue goes down , first they fuse with the nasal

    septum , because in the nasal cavity we have the nasal

    septum so all of these fuse together.

    The palate is completed by 60 days , the point of fusion is

    the point where the two sides meet with primary palate ,

    and then fusion goes in three directions .

    If we have a problem in fusion , this problem actually will

    be seen as failure of fusion , some people ( one case in

    every 700 birth ) are born with the cleft palate . why this

    happen ? It is because we have many reasons of fusion

    failure . most of the cases are because of genetic

    problems , so when the two palatine processes failed todevelop , they form what we call cleft palate , what

    happens when the primary palate fails to fuse with one or

    both of the secondary palates ? it gives cleft lip , the

    primary palate which also called the premaxilla is formed

    by the palatine processes of the maxilla so when the

    intermaxillary projections are failed to fuse to each other

    the resultant will be cleft lip and failure of the primary andsecondary palate to each other .

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    Sometimes when the patient is unlucky we will have a

    bilateral cleft . For example when the two palatine bones

    fail to fuse to each other and also fail to fuse with the

    primary palate ( premaxilla ) .These patients of cleft lip andpalate require multiple surgical procedures which needed

    to be completed before the age of 12 , so the

    management is going to be at early age .

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    Development of the maxilla :

    Just to remember we have two type of bone formation :

    0-Endochondral in which the preexisting cartilagetransform into bone

    2-Interamembranous no preexisting cartilageThe maxilla is formed by intramembranous

    ossification in which membranous mesenchymal

    tissue become bone.

    We have two ossification center in the maxillary

    development which are :

    0-Maxilla proper2-Premaxilla ( primary palate )

    Ossification at the maxilla proper begin 40 days

    after fertilization and become hollowed out later to

    form the maxillary sinus.

    The ossification of the maxillary proper begin

    below the infraorbital foramen from here the

    process of the maxilla arise which are :

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    0-Frontal process which fuse with themaxillary process of the frontal bone

    2-Palatine process

    3-Alveolar process which carry the teeth 4-Zygomatic processThe most important thing mentioned above is

    the medial and lateral alveolar plate which forms

    the alveolar process which carries the teeth

    Development of the mandible

    We will divide its development into two parts the

    body and the ramus

    The body is formed by intramembranous

    ossification , despite the fact we have cartilage

    in the region of the body of the mandible which

    called meckels cartilage but this cartilage have

    nothing to do with development itself it only

    guide the development so it act as scaffold

    The ramus of the mandible is formed by

    endochondoralossification , we have two site of

    cartilage : condylar cartilage : it appears at the

    11 week after fertilization and continues to

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    provide growth for the mandible until 21 years

    and it provide growth for the mandible in height

    Coronoid cartilage : it only active prenatally because just

    before birth it gets replaced by bone

    The other thing that we need to understand is that when

    we have muscle attachment there will be growth of bone

    The angle of the mandible had two muscles of mastication

    attached to which contribute to its growth :

    0-Medial pterygoid2-MasseterThe coronoid process has muscle attached to it

    which also contribute to its growth and called

    temporalis muscle

    Also in the mandible we have alveolar process which

    carry the teeth .

    Lecture 2 :Development of the tooth and its supporting

    structures

    The stomodeum (primitive mouth ) the primitive

    mouth is lined by ectoderm and beneath the

    ectoderm we have the mesoderm (and we said that

    mesoderm in this region the head region is notactually the real mesoderm it is ectomesenchymal

    tissue which means that it rises from neural crest

    cells ) the site where one tooth is going to develop

    we have condensation of ectomesenchymal tissue

    just under the ectoderm

    We start to see the condensation of the

    ectomesenchymal tissue and capillary networkbeneath oral epithelium at specific sites . Why at

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    specific sites ? Because we have more than one

    tooth and each tooth should be developed at specific

    site so at the site where the tooth should be

    developed we have condensation of theectomesenchyme

    Also we have capillary network at specific site then it

    leads to the formation of primary Epithelium band

    (PEB) when the oral epithelium thickens and

    invagenates into condensed ectomesenchyme . After

    that the PEB divides into two parts : one part goes

    buccully and also called vestibular lamina and theother part goes lingually and also called dental

    lamina

    The dental lamina is the structure that will develop

    the tooth and the vestibular lamina will become the

    sulcus (space ) it goes buccully and labially to form

    the oral vestibule ( vestibular lamina with time the

    cells inside it become lost and well get a space thisis the space between the teeth and the cheeks _

    posteriorly _ or the teeth and the lips _ anteriarly _ )

    and the remaining part which goes lingually is called

    the dental lamina

    So the vestibular lamina goes buccully to form the

    oral vestibule ( the space between the teeth and the

    cheeks or the teeth and the lips)The dental lamina goes lingually to form the teeth .Thats why its called arch shaped because we r

    talking about all teeth together _arch shaped banded

    tissue going lingually and it is surrounded by a

    condensation . At the end of each one it will form a

    teeth but not at the same time .A series of swelling

    happens at the deep surface , the terminal end of thedental lamina .These swelling at the terminal end is

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    called Enamel organ which is the swelling that

    happens in the first part of the tooth bud

    .

    This is a mandible this is the lower lip this is the area

    that is covered by oral epithelial as u see this is the

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    primary epithelial band (PEB) starts here and divided

    to vestibular lamina and lingual lamina.

    Lingual is a band as u see at the edge of this

    mandible we have swelling each swelling isresponsible of one of the teeth.

    Q: do u think that the band appear as one?? no each

    one will form a tooth but not at the same time as we

    said in dental anatomy for example mandible central

    incisor erupt before the maxillary central incisor

    We cant see all the tooth forming together

    crown formation

    *The dental lamina is responsible for formation ofprimary tooth but also each primary tooth in addition

    to the dental lamina which forms primary tooth there

    is another swelling responsible for forming the

    successor of that tooth ( successor tooth that come

    to replace the primary tooth )Q: what is the tooth successes the mandibular

    deciduous central incisor ?

    Permanent mandible central incisor

    That means if we lost the formed structure no

    permanent tooth will be developed if the deciduous

    tooth is missing also the permanent tooth ( the

    successor ) will be missing why ?? because both thedeciduous pre successor and the permanent

    successor they develop originally from the same

    germ layer

    But the opposite is not true if the primary is there but

    without swelling the permanent successor will not

    ***if the primary is missed no permanent will develop

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    What about non successor permanent teeth like

    permanent first, second and third molar???

    They will develop from this extension of the generallamina

    And here is the PEB (primary epithelium band )

    divide the vestibular lamina from dental lamina where

    we find swellings that will form the teeth

    Tooth germ : is the early part of the tooth that formed

    it includes the enamel organ and the surroundingectomesenchymle tissue and we divide it into 3

    different stages:0_bud stage

    2-cap stage3bell stage

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    Here is the first stage of development we call it bud

    stage it looks like a bud we see the oral epithelium

    invaginating against condensed ectomasnchyme

    later on this epethilium invaginating will start to forma concavity so it looks like a cap so we call it a cap

    stage then this cavity will become a big cavity and

    that will gives the bell shape then at the end of bell

    stage we start to see the formation of dentine in

    Dentinogenesis and the after a short while we start

    to see the formation of enamel which is

    Amelogenesis and finally the crown appear andcrown completion after this the root start to develop

    and that will lead to the eruption of the tooth :D

    Bud stage

    It involves the formation of enamel organ which will

    form the outer surface of the crown which is the

    enamel

    Enamel organ : early structure of tooth that comefrom ectoderm.

    So the enamel organ is the one which determine the

    three dimensional shape of the crown

    REMEMBER that teeth can't develop at the same

    time .It s impossible to see the entire tooth

    developing at the same time

    In the bud stage the enamel organ look spherical orovoid

    In the bud stage we cant differentiate the shape of

    the tooth it is poorly morpho differentiated so the

    three dimensional shape of the crown is not

    recognized and poorly histo differentiated so we cant

    recognize different type of cell .

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    Successive development of tooth germ involve

    complex interaction between epithelial and

    mesenchymal component

    In order to the development of the tooth there mustbe kind of interaction between the epithelium and

    ectomasynchyme or between ectoderm and

    ectomesnchyme so if we make tissue culture for the

    enamel organ there will be no tooth development

    because there no mesenchymal tissue which is

    needed for process to continue and the opposite is

    true and it is similar to the development of nails

    The basement membrane which separate between

    the epithelium and the underlying mesenchymal

    tissue play an important role in order to facilitate this

    interaction

    Remember the epithelium is the enamel organ and

    the epithelium is ectodermal origin .

    This is a tooth in the bud stage and here is the oral

    epithelium and underlying it we have the condensed

    ectomesenchymal tissue

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    Cap stage

    We divide it into early cap stage and late cap stageEarly cap stageThe down surface of enamel organ

    invaginate to form a cap shaped structureCell is still poorly differentiated : not similar to each

    other but still we able to differentiate between two

    populations of cell which are :

    1_ Inner ( central )_ related to the concavity _cellwhich have no specific function and it rounded in

    appearance

    peripheral cell which further divided into_2 (Internal enamel epithelium (IEE(External enamel epithelium ( EEE

    Late cap stage

    Here we have the development of cells called stellate

    reticulum which in the early cap stage were roundcell with no specific function but here in this stage the

    cell start to develop intercellular spaces.

    the IEE cell become columnar in shape

    the EEE become cuboidal

    the ectomesenchymal tissue which surround the

    enamel organ is recognized here in 2 different region

    1-Dental papilla which is the future pulp and dentine(related to the cells in the central of the concavity)

    2-Dental follicle is very important in the formation of

    the supporting structure of the tooth like cementum ,

    alveolar bone and periodontal ligament

    So the whole structure is called enamel organ it has

    a concavity that looks like a cap so it is called capstage of enamel organ at this stage we can identify

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    Then we have the bell stage which is divided into

    early and late

    What happens in early bell stage ?? The concavity

    deepens starts to become deep until the tooth lookslike a bell thats why it called the bell stage

    So what happens during the early bell stage???*Deferential cell division a long IEE ( not at the

    same way at different locations ) different rates of

    cell division at different sites that what gives different

    shapes of the tooth.

    What will happen if the rate of cell division rate is thesame in all sites?

    ***important***

    Mapping out the occlusal pattern of the crown*(Cessation of cell division ( stopping*

    For example if I have a cell division at a cusp or anincisal edge after awhile I dont want cell division to

    continue here because the tip of the tooth will swell

    *Active cell division At fissure sites and margin

    *Dental lamina breaks down

    *losing connection with oral epithelium

    *Dental lamina between tooth germs is lost

    -Remnants of dental lamina may remain as epithelialrests of series that may be involved in the aetiologyof cysts

    *We can identify here in early bell stage

    Histodifferentiation :

    We can identify different groups of cells each with

    different function

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    EEE*

    Cuboidal cells and they maintain the shape of the

    enamel organ and exchange substances betweenenamel and dental follicle

    **enamel organ is avascular in order to maintain the

    shape of the enamel

    Cervical loop*Increased cell division*

    At junction EEE and IEE*Stellate reticulum :are found in the center of enamel

    organ

    stellate because it is a star shape with many branchy

    processes and reticulum because it is a net work

    they have dendrites processes function in interacting

    nutrientsSR are those white cells

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    function : *it protects the underlying tissue against

    physical disturbance and that maintain the tooth

    shape

    its hydrostatic pressure is in equilibrium with that ofthe dental papilla allowing the proliferation of IEE to

    determine crown morphogenesis

    What happens if the hydrostatic pressure inside here

    in dental papilla area increase over the hydro static

    pressure of SR??The tooth will look like spherical ( it will swell )

    IEE : most important cells they are columnar

    cells that become elongated starting from the cusp

    tips and incisal edges

    Dental papilla : less differentiated than enamelorgan

    And now what happens In the late stage (

    appositional stage)Hard tissue formation ** dentine formation precedesenamel formation**

    The first part of the tooth that start to calcify is

    the tip of the tooth ( incisal edge or cusp tip)

    the last part of the crown that calcify is the most

    cervical partAppearance of a lingual down growth of EEE*

    In a deciduous tooth germs-Successor lamina

    -gives rise to tooth germs of permanent successorteeth

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    *In permanent tooth germs :it is a transient

    structure that disappears eventually

    Behind the second deciduous molars we should

    have first , second and third molars which are nonsuccessors dental lamina grows backwards to bud

    off permanent molars successively .

    :Reciprocal interactionvery important

    Dentine and Enamel formation commences at-Cusp tips

    -preameloblasts (mature IEE cells ) induce adjacentmesenchymal cells to become columnar and

    differentiate into odontobasts and secret predentine

    and dentine, then dentine induces ameloblasts to

    secrete enamel.

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    Done by :Ala'a Al_ Selawi Mais Maloul Al _Selawi

    o"It takes a little courage, and a little self-control, Ifyou want to reach the goal. It takes a great deal ofstriving, and a firm and stern-set chin. No matterwhat the battle, if you really want to win, there's noeasy path to glory. There is no road to fame. Life,however we may view it, Is no simple game; But itsprizes call for fighting For a rugged disposition thatwill not quit."

    GD LUCK TO U ALL