DEVELOPMENTAL BIOLOGY (LECTURE)

DEFINITION:
GAMETOGENESIS
• a discipline of biology concerned with the processes
and mechanisms that control and influence the
development and growth of organisms especially from
a molecular, cellular, or genetic perspective.
• Developmental biology of the embryo
• Embryo (Merriam Webster Definition)
- an animal in the early stages of growth and
differentiation that are characterized by cleavage,
the laying down of fundamental tissues, and the
formation of prim
- the young sporophyte of a seed plant usually
comprising a rudimentary plant with plumule,
radicle, and cotyledonsitive organs and organ
systems
MEIOSIS I AND II:
THE SPERM
 During the course of maturation, the sperm's haploid
nucleus becomes very streamlined and its DNA
becomes tightly compressed. In front of this
compressed haploid nucleus lies the acrosomal vesicle,
or acrosome.
 The acrosome is the cell’s Golgi apparatus and
contains enzymes that digest proteins and complex
sugars; thus, the acrosome can be considered a
modified secretory vesicle.
 acrosome and nucleus constitute the sperm head.
 An individual sperm is able to travel by whipping its
flagellum. The major motor portion of the flagellum is
the axoneme

PLANT GAMETOGENESIS:

THE AXONEME
 A structure formed by microtubules emanating from
the centriole at the base of the sperm nucleus.

MICROSPORANGIA: CONTAIN THE MICROSPORE

MOTHER CELL

DEVELOPMENT OF THE EMBRYO SAC:

THE EGG Summary of events leading leading to fusion
▪ The egg also accumulates a remarkable cytoplasmic
storehouse during its maturation
▪ Ribosomal proteins
▪ TRNA, ribosomes, mRNA, morphogenetic factors and
protective chemicals
▪ In most mammals the egg nucleus is still diploid—the
sperm enters before the egg's meiotic divisions are
completed.

Acrosome Reaction (initiated by the contact of sperm

with the egg jelly)
Contact causes the exocytosis of the sperm's acrosomal vesicle
and proteolytic enzymes and proteasomes (protein-digesting
EGG SURFACE complexes). Reaction of the sperm and the complex sugar of the
Vitelline membrane egg jelly (sulfate – containing polysaccharide the surface
receptors of the sperm membrane))

 Digestion of a path through the jelly coat to the egg

surface

 Sperm reaches the egg surface, the acrosomal process

adheres to the vitelline envelope and tethers the sperm
to the egg

 Proteasomes digest the vitelline envelope.

Binding with sperm receptors Initiates

 (1) a calcium transport channel that allows Ca2 to enter
the sperm head
Zona pellucida  (2) a sodium/hydrogen exchanger that pumps sodium
ions (Na+) into the sperm as it pumps hydrogen ions
(H*) out; and

 (3) a phospholipase enzyme that makes another second

messenger, the phospholipid inositol trisphosphate
(IP3, IP, is able to release Ca2 from inside the sperm,
probably from within

Acrosome reaction (2nd part)

 Extension of the acrosomal process.

 Polymerization of globular actin molecules into actin

filaments

 The influx of Ca2+ is thought to activate the protein

CELL CORTEX
RhoB in the
▪ Lying immediately beneath the cell membrane of most
is a thin layer (about 5 urn) of gel-like cytoplasm  acrosomal region and midpiece of sea urchin sperm
called the cortex.
▪ The cytoplasm in this region is stiffer than the internal  This GTP-binding protein helps organize the actin
cytoplasm and contains high concentration of globular cytoskeleton in many types of cells
actin molecules. During fertilization, these actin
molecules polymerize to form long cables of actin Role Of Bindin
known as microfilaments. Microfilaments are  Acrosomal protein mediating this recognition in sea
necessary for cell division. urchins is called bindin.
▪ They are also used to extend the surface into small
projections called microvilli.  Species-specific bindin receptors are found on the egg
▪ Membrane-bound, Golgi-derived structures contain vitelline envelope
proteolytic enzymes and are thus homologous to the
acrosomal vesicle of the sperm.
 Fusion of the Sperm and Egg Membranes
 The actin from the gametes forms a connection that
widens the cytoplasmic bridge between the egg and the
sperm.
 The sperm nucleus and tail pass through this bridge.
Fast block polyspermy
 Fast block to polyspermy is achieved by changing the
electric potential of the egg cell membrane. This
membrane provides a selective barrier between the egg
cytoplasm and the outside environment.
 Within 1-3 seconds after the binding of the first sperm,
the membrane potential shifts to a positive level, about
+20 mV due to the leakage of small amount of sodium
into the cell. Sperm can no longer attach with the
membrane with + potential.
 In normal monospermy, only one sperm enters the
egg, and a haploid sperm nucleus and a haploid egg
nucleus combine to form the diploid nucleus of the
fertilized egg (zygote).
 The sperm that enters the egg can provide a haploid
nucleus and a centriole to the egg.
 Polyspermy – entrance of numerous sperms leads to
disastrous consequences in most organisms
Slow block polyspermy
 Cortical granule reaction, a slower, mechanical block
to polyspermy that becomes active about a minute
after the first successful sperm-egg fusion
 Upon sperm entry the cortical granules fuse with the
egg cell membrane and release their contents into the
space between the cell membrane by exocytosis.
 Protein released:
 Trypsin-like protease called cortical granule serine
protease.
 This enzyme cleaves the protein posts that connect the
vitelline envelope proteins to the cell membrane, and
 it clips off the bindin receptors and any sperm attached
to them
 The components of the cortical granules bind to the
vitelline envelope to form a fertilization envelope
 This fertilization envelope is elevated from the cell
membrane by glycosaminoglycans released
 by the cortical granules.
 The fertilization envelope is then stabilized by
crosslinking adjacent proteins through egg-specific
peroxidase enzymes (soluble ovoperoxidase from the
cortical granules and Udxl in the former cortical
granule membrane) and a transglutaminase
Release of Calcium
- in sea urchins, it is thought that the binding all sperm to the
egg (or possibly the fusion of sperm and egg activates PLC7
through G proteins and Src kinases.
Fusion of the Gametes
 The mammalian sperm enters the oocyte while the
oocyte nucleus is "arrested" in metaphase of its second
meiotic division.
 The calcium oscillations brought about by sperm entry
inactivate MAP kinase and allow DNA synthesis.
 The mammalian oocyte still has chromosomes in the
middle of meiotic metaphase.
 Oscillations in the level of Ca2 activate another kinase
that leads to the proteolysis of cyclin (thus allowing
the cell cycle to continue, eventually resulting in a
haploid female pronucleus) and securin (the protein
holding the metaphase chromosomes together.
 DNA synthesis occurs separately in the male and
female pronuclei.
 The centrosome (new centriole) accompanying the
male pronucleus produces its asters (largely from
proteins stored in the oocyte)
 The microtubules join the two pronuclei and enable
them to migrate toward one another other
Fertilization
 Fertilization in sea urchin eggs occurs after the second
meiotic division, so there is a haploid female
pronucleus in the cytoplasm of the egg when the sperm
enters.
 The sperm's mitochondria and the flagellum
disintegrate inside the egg,
 Each gamete contributes a haploid genome to the
zygote\ the
 The mitochondrial genome is transmitted primarily by
the maternal parent
 Inside the egg, the sperm nucleus undergoes a
dramatic transformation as it decondenses to form the
haploid male pronucleus.
 A. First, the nuclear envelope vesiculates into small
packets, exposing the compact sperm chromatin to the
egg cytoplasm
 B. Proteins holding the sperm chromatin in its
condensed, inactive state are exchanged for other
proteins derived from the egg cytoplasm. This
exchange permits the decondensation of the sperm
chromatin.
  C. Once decondensed, the DNA adheres to the nuclear
envelope, where DNA polymerase can initiate
replication
 Decondensation is due to the phosphorylation of the of
the nuclear lamins and the sperm specific histones
 Done by cAMP dependent protein kinase.
 Phosphorylation changes the sperm histones to
histones in the egg cytoplasm.
 After the sea urchin sperm enters the egg cytoplasm,
the male pronucleus separates from the tail and rotates
180
 degrees so that the sperm centriole is between the
sperm pronucleus and the egg pronucleus.
 The sperm centriole then acts as a microtubule
organizing center, extending its own microtubules and
integrating them with egg microtubules to form an
aster.
 Microtubules extend throughout the egg and contact
the female pronucleus, at which point the two
pronuclei migrate toward each other.
 Their fusion forms the diploid zygote nucleus
Mammalian Fertilization (Internal)
 Translocation
 Ampulla
 Region of the oviduct where fertilization
takes place
 Egg
TRANSLOCATION
 Cumulus oophorus
 Very important for the fimbriae of the oviduct to
capture the egg.
 Once it is picked up, a combination of ciliary beating
and muscle contractions transport the oocyte-cumulus
complex to the appropriate position for its
fertilization in the oviduct.
 Sperms are transported to the oviduct mainly by the
muscular activity of the uterus.
 Sperm (flagellar) motility is important once sperm
arrive within the oviduct; sperm become hyperactive
in the vicinity of the oocyte.
 Directional cues
 Temperature and chemical
 From the vagina to the ampulla the sperm matures.
CAPACITATION
The set of physiological changes by which sperm become
competent to fertilize the egg is called capacitation.
The sperm cell membrane is altered by the removal of
cholesterol by albumin proteins in the female reproductive
tract- changes the location of the lipid rafts that contain the
receptors and clusters in the anterior sperm head.
The membrane potential of the sperm cell membrane becomes
more negative as potassium ions leave the sperm- calcium
channels open: activation of the cAMP
Protein phosphorylation. Heat-shock proteins migrate to the
surface of the sperm. Here, they may play an essential role in
forming the receptor that binds to the zona pellucida.
The outer acrosomal membrane changes and comes into contact
with the sperm cell membrane in a way that prepares it for
fusion.
Hyperactivation, Thermotaxis and Chemotaxis
 Hyperactivation, along with a hyaluronidase enzyme
on the outside of the sperm cell membrane, enables the
sperm to digest a path through the extracellular matrix
of the cumulus cells .
 Sperm moves from cooler to hotter part
 Cunulus oophorus cells secrete chemicals (i.e.
progesterone) that attract the sperm
Recognition of the zona pellucida
 The first step appears to be a relatively weak binding
accomplished bv the recognition of a sperm protein by
a peripheral protein that coats the zona pellucida.
 This is followed by a somewhat stronger association
between the zona and the sperm's SED1 protein.
 Last, a protein on the sperm (and possibly several other
factors) forms strong links with the ZP3 of the zona.
 Acrosome reaction
Gamete Adhesion
 initial tethering is accomplished. It appears that sperm
initially bind to a 250-kDa protein that is associated
with, but not integrally part of, the zona pellucida
 Sperm binding is also facilitated by the sperm
adhesion protein SED1, which binds to the zona
protein complex
 Binding to the ZP3 glycoproteins.
 Some of these sperm proteins bind to the serine- and
threonine-linked carbohydrate chains of ZP3, and one
of the zona-binding proteins, a sperm-surface
galactosyltransferase, recognizes carbohydrate
residues on ZP3
 Another set of proteins that bind to the zona are
ADAM3 and ADAM2 (also called cyritestin and
fertilin p, respectively).

 ZP3 also initiates the acrosome reaction after sperm

 ZP3 is able to cause the Ca_' -mediated exocytosis of

the acrosomal vesicle.

 ZP3 binding, phospholipase C (which synthesizes IP-)

is activated, the sperm head becomes more alkaline,
and membrane Ca2" channels are opened.

 ZP3 initiates these events by crosslinking the sperm

cell surface galactosyltransferases, whose active site
faces outward and binds to the carbohydrate residues
of ZP3

 This crosslinking activates specific G proteins in the

sperm cell membrane, initiating a cascade that opens
the membrane's Ca2+channels and results in the
calcium-mediated exocytosis of the acrosomal vesicle

 Exocytosis release of proteases and lysis of the ZP and

creates a hole for which the sperm can travel into the
egg.

 zonadhesin protein from the acrosome may fix the

sperm to the point of attachment and provide a pivot
point for the sperm to enter into the zona

 Second, certain proteins in the inner acrosomal

membrane specifically to the ZP2 glycoprotein

 Third, in some species, proacrosin, a protein that

adheres to the inner acrosomal membrane, binds to
sulfated carbohydrate groups on the zona pellucida
glycoproteins

 In mammals, the sperm contacts the egg not at its tip

(as in the case of sea urchins), but on the side of the
sperm head .

 . The junction between this inner acrosomal membrane

and the sperm cell membrane is called the equatorial
region, and this is where membrane fusion between
sperm and egg begins

 In sea urchin gamete fusion, the sperm is bound to

regions of the egg where actin polymerizes to extend
microvilli to the sperm

 Mammalian gamete fusion may depend on interaction

between a sperm protein and integrin-associated CD9
 EMBRYOGENESIS
Morphogenesis – establishment of the plant body.
Division of the zygote – Single terminal cell embryo
- The zygote divides by mitosis forming the smaller
apical/terminal and larger basal cells.
- The start of the embryo. The apical cell produces the embryo
proper.
- a basal cell, and a suspensor cell.
Dermatogen Embryo
- each octant has undergone periclinal division
- formation of a protoderm (first recognizable tissue
differentiation at the outermost layer)
- The outer cells perform anticlinal division to form the
protoderm

Two terminal Cell Embryo 2-cell stage to Dermatogen Embryo

- Single apical cell divides by mitosis.
- Basal Cell (BC) forming the suspensor and at least part of the
root.
Quadrant Embryo
- Longitudinal divisions in the two terminal cells will produce
a four-celled embryo-proper.
- The suspensor now consists of three cells including the basal
cell.
HORMONAL CONTROL
The basis of many cell types is established, through the process
of axis and pattern formation.
Both the apicobasal and radial axes are formed at this time.
Pattern formation in the embryo strongly depend on the
activity of the hormone auxin.
Export of auxin occurs actively through the PIN-FORMED
protein family of auxin exporters.

Octant stage Embryo

Three rounds of cell division:

1. First two rounds of longitudinal divisions at right

angles to form four cells of equal size.

2. One transverse division that gives two tiers, producing

an eight-cell stage also called the octant stage embryo

At the octant stage, the apical–basal axis of the embryo can be

distinguished as the apical/upper tier embryo domain (UT),
which generates the shoot apical meristem (SAM) and

Most of the cotyledons, and the basal/lower tier embryo domain

(LT), which will contribute to the hypocotyl, RAM, and parts
of the cotyledons.

The BC divides transversely, generating a filament of seven to

nine cells by the globular stage of EP development

• giving rise to the extraembryonic suspensor (S).

TAKE NOTE!
Genes:
yoda (yda), gnom (gn), grounded (grd), short suspensor
(ssp), wrky2
The YDA pathway is involved in zygote elongation and
development of basal cell lineages in Arabidopsis

Genetic Control:
WUSCHEL:
• To regulate the maintenance of the shoot apical
meristem is the
• Family of WUCHEL RELATED AUXIN IN EMBRYOGENESIS
HOMEOBOX:
• ARF (Auxin Response Factors) gene expression
• WOX GENES:
attributed to the embryo specific auxin responses in
The role of WOX genes in early embryogenesis becomes
different cell types.
most apparent in the interplay between the WOX2 and
• As different cell types
WOX8/9 genes
• all have a distinct set of ARFs, they will
WOX2 and WOX8/9 genes are initially, coexpressed in the
respond differently to the same stimulus,
undivided zygote but they become restricted to the apical
which will result in a different
(WOX2) and basal (WOX8/9) cell after the first division.
developmental output.
Initiated Upon Fertilization:
YDA responsible for the kinase cascade
YDA encodes a mitogen-activated protein kinase
(MAPKKK).
MAPKKKs participate in signaling events via aMAP kinase
cascade, involving MAPKKs (MKK4 and 5 in the YDA
pathway)
WRKY DNA-BINDING PROTEIN 2 (WRKY2) was shown
to be a target of the YDA/MKK4/5/
MPK3/6 kinase cascade
 Early Globular Embryo
- The protoderm and inner cells of the embryo-proper divide by
anticlinal and longitudinal divisions, respectively
- All cells are interconnected by many plasmodesmata
- Division within the suspensor is complete by this stage
including the hypophyseal and basal cells.
- consists of seven to nine (eight in this example) highly
vacuolate cells.
Late Globular Embryo
- The suspensor is very elongated (completed its divisions)
- The micropylar chamber of the embryo sac contains a large
volume of free-nuclear endosperm which surrounds virtually
the whole of the suspensor
- the hypophysis, derived from the top-most cell of the basal cell
lineage, divides asymmetrically to form the precursor of the
quiescent center (QC), which is a smaller lens-shaped cell, and
another larger basal cell which is a precursor of the distal stem
cells of the root meristem, the columella
Molecular Control – Globular Stage Embryo
Stem cell and tissue specification
• AUXIN RESPONSE FACTOR5/ MONOPTEROS
(MP) gene:
• key regulator in hypophysis specification and
specification of other cells that form the embryonic
root
• TARGETOFMONOPTEROS(TMO) genes.
• TMO5 and TMO7 encode basic helix–loop–
helix (bHLH)transcription factors and both
are only transcribed in the basal inner cells of
the proembryo
• TMO5
• form a dimer with another member of the
bHLH family, LONESOME HIGHWAY.
• For periclinal division of the vascular tissue.
• TMO7 protein was shown to move from its
transcribed region to the hypophysis.
The PLETHORA (PLT) transcription factors
• PLT1 and PLT2 are key regulators of root meristem
maintenance and root ‘fate’
The Shoot Meristem
• Exclusion of PLT gene expression from the apical
embryo domain is required for shoot formation
• CLASS III HOMEODOMAIN-LEUCINE ZIPPER
genes (HD-ZIP III – inhibits the PLT
• expressed in the apical part of the embryo and
regulated by MICRO-RNA165/166
Heart-shaped Embryo
- Embryo assumes a triangular profile.
- Cells of the procambium and shoot apex are more vacuolate
than those of the protoderm.
- Cotyledon initials are present and the central cells are
beginning to elongate and divide to form the provascular tissue
- Uppermost cell of the suspensor has started to differentiate to
form the hypophysis (h).
Torpedo Stage Embryo
- The cotyledons are elongate by this stage, having forced their
way into the fully cellularized endosperm.
- The central cells of the radicle and hypocotyl have
differentiated, forming the provascular tissue.
--------------THE SAM and RAM is established----------------
- Small populations of cells within the SAM and RAM divide
slowly to provide the surrounding stem cells, and these regions
are called the organizing centers.
------------The Vegetative Shoot Apical Meristem--------------
- The vegetative shoot apical meristem consists of a small group
of undifferentiated cells located at the distal end of the shoot,
and is surrounded by the youngest leaf primordia.
------------The Shoot Apical Meristems (SAM)------------------
- The shoot apical meristem gives rise to the entire above
ground portion of the plant except the hypocotyl and cotyledons.
- The shoot apical meristem has several functions:
- it initiates tissues,
- initiates organs, communicates signals to other
parts of the plant, and
- maintains itself as a formative region
----------------------------Juvenile SAM-----------------------------
- The primordia for leaves 1 and 2 have enlarged and adopt a
dorsiventral symmetry.
- Trichomes become conspicuous on the adaxial surface at the
distal end of each primordium.
- Stipules have enlarged and begun to differentiate into club-
like structures. The meristem is still flat.
-----------------------------Mature SAM-----------------------------
- The vegetative shoot apical meristem changes from a
relatively flattened, bilaterally symmetric structure to a dome
shaped, radially symmetric structure consisting of
approximately 450 cells.
SHOOT APICAL MERISTEM:
Two principal populations of cells:
1. The tunica, constituting the surface mantle of cells
undergoing anticlinal divisions
2. Inner corpus cells dividing in all planes. Layers:
• LI, (dermatogen)
• L2 (periblem)
• L3 (plerome).
The Central Zone and Organizing Centers:
- The central zone (CZ) contains relatively slowly dividing cells
that are slightly larger and more vacuolated than the
surrounding cells. It harbors the stem cell niche, consisting of
the stem cells in the outermost three cell layers and the signaling
niche cells, termed the organizing center (OC).
- OC maintains the stem cells in an undifferentiated state.
- Stem cells express CLAVATA 3 (CLV 3): ligand for receptor
kinase and restrict WUS expression and limits the size of the
OC.

Outgrowth of Organ Primordia: Protoderm, cotyledon and leaf
primordia
• CZ unvacuolated and divide more rapidly and, based
on gene expression, initiate differentiation.
• SHOOTMERISTEMLESS (STM) - Allow
the stem cells to amplify.
• STM becomes downregulated in the organ
precursor cells and the expression of organ-
specific genes is initiated, resulting in the
outgrowth of organ primordia from the flanks
of the meristem.
• The differentiation of the outer layer and
inner cells accords with the gene expression
pattern of two homeodomain
• PROTODERMAL FACTOR 2.
• initially detected throughout the
early EP, but immediately after the
tangential divisions, expression
becomes preferential to the
protodermal layer
• WOX2 for protoderm formation
• The CUP-SHAPED
• COTYLEDON (CUC) NAC-domain transcription
factor genes CUC1, CUC2, and CUC3 are specifically
expressed in boundary positions
• The expression of AINTEGUMENTA,
• Cotyledon formation from the general
embryonic tissue.
• Leaves initiate from PZ cells that are amenable to
differentiation.
• Auxin plays a key role in specifying those
cells, as only the PZ cells overlapping
periodic auxin maxima are designated to
develop into leaves.
The Procambium
• Start at the 4 initial cells during the globular stage.
• After their specification, these cells elongate and
undergo oriented and coordinated cell divisions,
thereby increasing the number of procambial cells and
establishing their typical strand-like anatomy up to the
mature embryo stage
• a subset of procambial cells undergoes asymmetric
divisions, generating precursors of phloem and xylem
cells while maintaining a pool of procambium cells
between these tissues.
• Auxin-dependent transcription factor
MONOPTEROS/AUXIN RESPONSE FACTOR 5
(MP/ARF5) plays a major role in translating auxin
accumulation into the establishment of procambium
identity.
• Helix-loophelix (bHLH) transcription factor
TARGET OF MONOPTEROS5 (TMO5) is first
expressed in all four procambium initials at the
globular stage
The Root Apical Meristems (RAM)
- an amalgam of cell populations of two different clonal origins.
The root cap, quiescent center, and part of the root cortex arise
by divisions of the suspensor cell closest to the embryo, known
as the hypophysis; the rest of the embryonic root is derived from
cells cut off by the apical cell.
Composed of four sets of initials, or stem cells.
- Periclinal divisions of a distal plate of initial cells (columella-
root cap initials)- give rise to the group of cells constituting the
columella.
- A ring of about 16 initial cells (epidermal-lateral root cap
initials) generates by periclinal divisions the lateral root cap
cells, which envelop the columella cells. Epidermal cells of the
root are also derived from these cells.
- The cortical and endodermal cells can be traced to the middle
layer of initials (cortex-endodermal initials) that undergo
periclinal divisions.
The cells of the vascular cylinder including the pericycle are
derived from a proximal plate of initials (pericycle—vascular
initials).
Pericycle
The presumptive founder cells of primordia are derived after the
formation of short initial cells by anticlinal divisions
The Primary Meristems
• The differentiation of the outer layer and inner cells
with the gene expression pattern of two homeodomain
• leucine zipper class IV (HD-ZIP IV) TFs :
• 1. ARABIDOPSIS THALIANA MERISTEM
LAYER 1 (ATML1)
• 2. PROTODERMAL FACTOR 2 (PDF2)
Ground Meristem
• MP, that ‘marks’ the inner cells at the dermatogen
stage, has a role in ground tissue initiation and
provascular meristem development.
• The ground tissue daughter cells will then divide to
give the endodermis and other cells of the cortex;
• Auxin activity was found to be highest in vascular
tissue, lowest in the protoderm, and intermediate in the
ground tissue in globular stage embryos.
Derivatives of the Apical Meristems
• The derivatives of the Apical meristems persist in the
floral meristems.
The Floral Meristems
FLC (flowering Locus)
SOC (Supressor of overexpression of Constant 1)
 Lateral Meristems
The floral meristems derived from the L1,L2,L3
• Flank meristem (2)
• File meristem (3)
• Central initiation zone (1)

Control from leaf to floral formation Fascicular and interfascicular meristems

The formation of the interfascicular meristems starts when the

parenchymous and/or endodermal cells located to the both sides
of the fascicular cambial cells begin to divide

Ground meristems: Derivatives of the corpus

The ground tissue is specified at early globular stage of the
embryo.
In the apical meristems, the layer of cells known at the corpus
divide in any plane to form cells to become the ground meristem.
Periclinal divisions of the ground tissue daughter cells generate
separate endodermis and cortex layer.

Pattern of the Floral Whorls: Examine the concentric pattern of

the emergence of the floral whorls
 Cork Cambium (secondary growth) SOMATIC EMBROGENESIS

Ongoing secondary growth within the root stele forces the non-
dividing outer cell layers to peel off. Thereafter, the root's outer
protective layer is replaced by a barrier of cork cells (phellem)
produced by a lateral meristem called cork cambium
(phellogen), which originates from the pericycle. The cork
cambium also produces parenchyma (phelloderm) cells inwards.
Collectively, these cell types are called periderm.

The Complete Embryo in Seed (Seedling)

Hypophysis: upper most suspensor cell recruited to the embryo.
Derivatives:
Columella
Quiescent center

Origin of the Epicotyl, Plumule and Shoot

The terminal cell forms the embryo proper, and by octant
stage has generated an upper tier and lower tier of cells. The
upper tier is fated to form the SAM and bulk of the
cotyledon.

The Origin of Radicle and the Hypocotyl:

By the triangular stage of embryogenesis, the lower tier
divides into the upper lower tier (ult) and the lower lower tier
(llt), the latter giving rise to all cells of the radicle and the
hypocotyl.