Houari Boumediene University of Science and Technology

(USTHB)
Faculty of Biological Sciences (FSB)

First year Licence -SNV /2023-2024

Animal Biology subject

General embryology course summary

Gametes
Fertilisation
Segmentation
Gastrulation

Collective of Biology teachers L1-SNV-FSB-USTHB

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 Gametes
I - Notion of soma and germen

Sexual reproduction is made possible by cells specialised in reproduction, or gametes.

Gametes are haploid cells, with n chromosomes; they carry half of the paternal or maternal
genetic message.
These gametes are produced in specialised organs, the gonads, the male gonad or testis (poly
p.9, fig.1); the female gonad or ovary (poly p.17, fig.1A) from a particular category of cells,
the gonocytes or primordial germ cells. These primordial germ cells separate very early from
the three embryonic layers.
The three embryonic layers are at the origin of the SOMA or set of body cells known as
somatic cells.

The gonocytes are the origin of the germ cells that evolve to form haploid gametes. All the
gonocytes and their descendants constitute the GERMEN.

II - The male gamete (spermatozoon)

1-The stages in its formation or spermatogenesis (see OPU book, 2017, p.13, fig.6).
1.1 Multiplication: This occurs continuously from puberty (passage to adulthood with the
ability to reproduce). The cells that undergo these mitoses (maintenance of the 2n number of
chromosomes) are spermatogonia. Strain spermatogonia (with a dark dusty nucleus) always
give a strain spermatogonia and a pale dusty spermatogonia which divides by successive
mitoses into crusty spermatogonia. The exact number of mitoses varies from species to
species.
1.2 Growth: cytoplasmic growth is observed in crusty spermatogonia which become
auxocytes called spermatocytes I. The 2n number of chromosomes is therefore retained.

1.3 Meiotic maturation : Each spermatocyte I undergoes the first reductional division of
meiosis and gives two spermatocytes II (with n fissured chromosomes and 2q DNA) which
undergo the second equational division of meiosis, each giving two round spermatids with n
chromosomes and q DNA.

1.4 Cytoplasmic differentiation into spermatozoa or spermiogenesis : the spermatid

transforms into a spermatozoon (see OPU book, 2017, p.14, fig.7).

2-Ultrastructure of the spermatozoon (see OPU book, 2017, p.15, fig.8)

The spermatozoon is constitueted by 3 parts:
-Head: it contains the nucleus and the acrosome.
-Neck: it contains the proximal centriol, distal centriol and segmented columns.
-Flagellum: formed by parts, the intermediate part, the main part (the tail) and the terminal
part.

3 -Characteristics of the male gamete

-It is a small cell with no cytoplasm.
-It is produced in large numbers and continuously.

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- It carries the paternal genetic message in n-chromosome nucleus.
-It is mobile (after remaining in the epididymis) and therefore able to move towards the
female intervention of microtubules and mitochondria of the flagellum.
-It is capable of lysing the envelopes of the female gamete by the enzymes contained in
acrosome.

4 -The somatic cells of the testis (see OPU book, 2017, p.11, fig.5)
In the testis, inside the seminiferous tubules, alongside the germ cells, somatic cells can be
seen, the Sertoli cells which play a nourishing and hormonal role. The interstitial tissue
contains Leydig cells, which produced hormones, mainly testosterone.

III. The female gamete

1-The stages of its formation or oogenesis (see OPU book, 2017, p.21, fig.3).

1.1. Multiplication: oogonia multiply during embryonic life.

1.2. Oocyte growth: takes place in several stages: it begins during embryonic life: the ovary
at birth does not contain oogonia but primary oocytes (oocytes I). It continues during
childhood.
This oocyte growth will resume at each cycle for some primary oocytes.
For eggs that contain reserves, it is during this period that these reserves are accumulated. For
all eggs this will be the time when maternal ribosomes are accumulated.

1.3. Nuclear or meiotic maturation (meiosis) takes place over several stages.
During fetal life, prophase of the first division of meiosis is initiated and then stopped.
In vertebrates, during the hours preceding ovulation, this first division is completed and
produces an oocyte II. The second division of meiosis will only take place if fertilization is
achieved.
On the contrary, in the sea urchin, the two divisions of meiosis are completed before
fertilization.
Cytoplasmic division is unequal and the cytoplasm is retained in a single ovotid or ovule with
formation of polar globules.

2-Different types of eggs (see OPU book, 2017, p.24, fig.5; p.27, Table 1).
The presence or absence of yolk makes it possible to distinguish:
- Eggs completely devoid of yolk: aleciths example: Mammals
- Eggs with little yolk: oligoleciths example: Sea urchins
- Eggs with an average quantity of yolk: mesoleciths. This yolk is arranged heterogeneously
in a yolk gradient: heteroleciths example: Amphibians
- Eggs with very abundant yolk: megaleciths where the yolk occupies a central place:
centroleciths example: Insects or almost total: teloleciths example: Birds.

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3- The egg envelopes and accompanying cells
Around the plasma membrane, a vitelline membrane (in yolk eggs) (see OPU book, 2017,
p.25, 26, fig.6a-c) or a pellucida zone (in alecith eggs) of a glycoprotein nature develops (see
OPU book, 2017, p.26, fig.6 bis d). It contains the specific sites of the spermatozoon. A
mucus gangue often envelops the vitelline membrane and swells on contact with water (sea
urchins, amphibians) (see OPU book, 2017, p.25, fig.6a).
In birds, the albumen or egg bank is deposited around the yolk (representing the female
gamete itself). A shell membrane and a shell envelope the whole (see OPU book, 2017, p.25,
fig.6b).

In mammals, the follicular cells accompanying oocyte I will evolve and take part in the
formation of follicles (see OPU book, 2017, p.26, fig.6 bis d). Some of these will accompany
oocyte II at the time of egg laying or release of the gamete into the oviduct.

4-Characteristics of the female gamete

Large cell.
Large cytoplasm rich in ribosomes, will be the cytoplasm of the fertilized egg.
Produced cyclically and in limited numbers (atresia phenomenon).
Nucleus with n chromosome (maternal genome).
Immobile cell, leads a slowed life.

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 Fertilisation
I-Definition
This is the starting point of embryonic development, the union of the male gamete and the
female gamete. A zygote or fertilised egg is formed by the return to diploidy.

II-Methods of fertilisation
1-External fertilisation: the gametes are released outside the genital tract and are found in
the water (sea urchins, amphibians).
2-Internal fertilisation: the gametes meet in the female genital tract (birds, mammals).

III-Gamete encounters
1- Gamete pathway: in species with external fertilisation (sea urchins), the spermatozoa are
attracted to ovules and agglutinate (action of the fertilisins released by the ova - LILLIE
experiment, 1912); Spermatozoa are recognised by eggs of the same species.
In species with internal fertilisation (birds, mammals), the meeting takes place in the female
genital tract.

2- Capacitation: before fertilising the female gamete, the spermatozoa undergo the
phenomenon of capacitation, which corresponds to the removal of inhibition of the acrosome
enzymes. This takes place in the gangue (sea urchin, amphibian) or the cumulus oophorus
(human). Capacitation consists of various modifications (membrane constituents,
metabolism, sperm motility).

IV- Main stages of fertilisation in mammals

1-Recognition, adhesion/fixation
1-1- Adhesion of the spermatozoon to the zona pellucida (OPU book, 2017, p.34, fig. 5)
1-2- Acrosomal reaction (OPU book, 2017, p.34, fig. 6)
1-3- Penetration through the zona pellucida (OPU book, 2017, p.35, fig.7)
2-Fusion of gametes (OPU book, 2017, p.36, fig.9A, B, C): plasmogamy

V- Consequences of fertilization
1-Activation of the egg
1-1-Blocking of polyspermy
In all vertebrates, polyspermia is blocked (except in birds).
1-1-1- Early blockage: change in membrane potential (Na+ input).
1-1-2-Late blockage: Cortical reaction (OPU book, 2017, p.38, fig.10A, B), this is the
release by exocytosis of the contents of the cortical granules which leads to:
a- Modification of sperm receptor sites by enzymes (supernumerary spermatozoa detach).
b- Detachment of the vitelline membrane from the plasma membrane (rupture of the binding
proteins by enzymes contained in the cortical granules).
c- The penetration of water between the two membranes (osmosis) and the formation of a
perivitelline space (action of proteoglycans).

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d- The deposition of structural proteins against the plasma membrane forms the hyaline layer
which, by thickening, helps to maintain the blastomeres during segmentation.
e- The deposition of structural proteins at the vitelline membrane forms the fertilisation
membrane which is impermeable to spermatozoa.

1-2-Completion of meiosis
1-3-Amphimixis (OPU book, 2017, p.39, fig.11)
-Spermaster or monoaster stage
-Amphiaster stage
-Diaster stage
-Amphimixis stage

2-Structural changes in amphibians

2-1-Orientation (or equilibration) rotation (OPU book, 2017, p.40, fig.13)
This is due to the detachment of the two plasma and vitelline membranes; the free egg in the
perivitelline space tilts and is oriented with the animal pole upwards and the vegetative
(heavier) pole downwards. This equilibrium position is determined by gravity.

2-2- Symmetrisation rotation (OPU book, 2017, p.41, fig.14)

In the region opposite the point of penetration of the spermatozoon, a light zone appears due
to the sliding of the superficial cortical layer which marks the future dorsal surface: this is the
grey crescent. The plane passing through the PA-PV axis and the point of penetration of the
spermatozoon cuts the grey crescent into two equal halves: this is the plane of bilateral
symmetry.

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 Segmentation

I. General Overview
Segmentation allows the formation of a multi cellular organism from a single cell
(zygote). It corresponds to successive mitoses of the zygote into daughter cells or
blastomeres without an increase in total volume or modification of the external shape of
the germ, which passes through a morula stage (resembling a small blackberry), and
then develops an internal cavity, the blastocoel (or blastocystic cavity in humans). This
segmentation cavity is formed either by cellular spacing (amphibians, mammals) or by
yolk lysis (birds). The germ obtained at the end of the segmentation is called blastula or
blastocyst (human).

II. Different Modes of Segmentation

There are several modes of segmentation depending on the richness of eggs in yolk and
its distribution. The presence of yolk (yolk plates) prevents division or cleavage planes.
Two main types of segmentation are distinguished:

1-Total or Holoblastic Segmentation

Cleavage planes affect the entire fertilized egg. This type of segmentation is observed
in oligolecithal, heterolecithal, and alecithal eggs. Segmentation can be equal
(identical blastomeres), subequal (almost equal blastomeres), or unequal (blastomeres
of different sizes), which leads to the formation of macromeres, micromeres, and
sometimes mesomeres. Segmentation can be synchronous, where cell divisions occur
at the same rate, or asynchronous, where blastomeres divide at different rates
(e.g.,micromeres divide more rapidly than macromeres).

2-Partial or Meroblastic Segmentation

Only a portion of the egg cytoplasm, characterized by its low yolk content, undergoes
division. This occurs in megalecithal telolecithal and centrolecithal eggs. Two
different modalities are observed:
 Partial discoidal segmentation: e.g.,birds (only the germinal disc divides).
 Partial superficial or peripheral segmentation: e.g.,insects (only the egg's
periphery divides).

III. Comparison of Segmentation in Certain Vertebrates

 Amphibians: Holoblastic or total, unequal, and asynchronous segmentation (see OPU
book, p.48, fig.2).
 Birds: Meroblastic or discoidal partial, unequal, and asynchronous.
Mammals (Humans): Holoblastic or total, subequal, and somewhat synchronous (see
OPU book, p.51, fig.5).

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 Gastrulation in Amphibians
I-Definitions and methods of study

1-Definitions
Gastrulation ensures the establishment of the three embryonic layers: ectoderm, mesoderm
and endoderm thanks to Morphogenetic movements.
The internal penetration movement is Embolia.
The movement of extension on the surface is Epibolia.
The winding movement on a treadmill is the movement of Involution.
When the territory spreads out, there is Divergence.
When the territory is grouped into a smaller volume, there is Convergence.

2-Methods of study
It is the tattooing of the germ with vital colorings which made it possible to follow the fate of
every zone (see OPU book, 2017, p.54, fig.1) and made it possible to establish the map of the
presumptious territories of the blastula (see OPU book, 2017, p.55, fig.2).
II-Morphological phenomena

1- Dorsal external views

Below the gray crescent, invagination first forms the dorsal lip of the blastopore (see OPU
book, 2017, p.56, fig.3A).
By acquiring lateral lips, this blastopore forms an arc or horseshoe (see OPU book, 2017,
p.56, fig.3B); it then becomes circular by the formation of a ventral lip (see OPU book, 2017,
p.56, fig.3C, D).
At the end of gastrulation which lasts 24 hours, the diameter of the blastopore gradually
reduces (see OPU book, 2017, p.56, fig.3E).

This blastopore is continually crowded with macromeres rich in yolk or yolk cells of the
endoblast.

III- Analysis of movements of presumptive territories

1- Outer layer
It gradually spreads over the surface and compensates for the penetration of other territories.
It undergoes an epiboly movement (see OPU book, 2017, p.59, fig.7A-E). At the end of
gastrulation, the germ shows us a dorsal neurectoblast and a ventral epiblast, constituting the
outer envelope of the gastrula (see OPU book, 2017, p.59, fig.7E).

2- Inner layer
It penetrates en masse like a plug that is pushed into the blastopore: it undergoes an embolism
movement. It constitutes the floor of the new cavity which is called Archenteron (ancestor of
the digestive tract). It pushes back the old blastocoel which gradually decreases in volume and
disappears at the end of gastrulation (see OPU book, 2017, p.57, fig.5A-E). Finally, formed
of cells rich in yolk, it causes the rocking movement of the germ which places its heavy belly
(rich in yolk) downwards.

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 3- Pharyngeal endoblast located above the dorsal lip
This small territory enters first, as soon as the notch is formed, by wrapping around the dorsal
lip and internally lining the territory remaining outside (see OPU book, 2017, p.57, fig.5A-
E).

4- Precordal and cordal territories

They undergo the treadmill winding movement (involution) around the dorsal lip of the
blastopore, following the pharyngeal endoblast. These territories converge and are placed in a
mid-dorsal position against the internal face of the dorsal neurectoblast remaining outside and
form the roof of the new cavity or archenteron (see OPU book, 2017, p.57, fig.5A-E).

5- Mesoblast
It converges towards the lateral then ventral and finally dorsal lips of the blastopore, rolls up
by involution and spreads inside the germ to the left and right of the cordal territory to which
it remains attached.
It is in cross section that the establishment of this mesoblast can be observed. we note that
initially attached to the endoblast, the mesoblast subsequently separates from it and then
becomes a true middle layer or mesoderm. It then progresses towards the ventral side (see
OPU book, 2017, p.60, fig.8A-C).

Conclusion
In approximately 24 hours, the germ became a gastrula made up of three layers; ectoderm
(outer layer), mesoderm (middle layer), and endoderm (inner layer).
Bilateral symmetry, established by the formation of the gray crescent, is morphologically
visible at the end of gastrulation because the germ tends to stretch in the antero-posterior
direction.

For more information:

COLLECTIF D’ENSEIGNANTS DE BIOLOGIE L1-SNV de L’USTHB, 2017- Cours
illustré de Biologie Animale, Tome 1 Embryologie générale, Edition OPU, Alger, 99p.

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 Translation of French/English figures legends
French English
1er globule polaire 1st polar globule
1er globule polaire (n) GP1 1st polar cell (n) GP1
2ème globule polaire (n) GP2 2nd polar cell (n) GP2
Acrosine Acrosine
Acrosome Acrosome
Albumen Albumen
Annulus Annulus
Appareil de Golgi Golgi apparatus
Axonème Axoneme
Blastocœle Blastocoel
Bouton embryonnaire Embryounicbutton
Cavite Blastocystique Blastocysticcavity
Cellule de Leydig Leydigcell
Cellule de Sertoli Sertolicell
Cellule souche:ovogonie (2n) Stem cell: ovogony (2n)
Centriole distal Distal centriole
Centriole proximal Proximal centriole
Centrosome Centrosome
Chalaze Chalaza
Chambre à air Air chamber
Coiffe post-acrosomienne Post-acrosomal cap
Colonne segmentée Segmented column
Condensation de la chromatine (C) Chromatin condensation (C)
Coquille Shell
Corona radiata Corona radiata
Couche enveloppante ou trophoblaste Wrap-around layer
Couche hyaline Hyaline layer
Coupes transversales Cross-sections
Crêtes neurales Neural crests
Croissant gris Grey crescent
Cytoplasme Cytoplasm
Cytoplasme résiduel Residualcytoplasm
Disque germinatif (cytoplasme+noyau) Germ disc (cytoplasm + nucleus)
Division équationnelle Equational division
Division réductionnelle Reduction division
Doublet de microtubules Microtubule doublet
Endoblaste non pharyngien Non-pharyngeal endoblast
Endoblaste pharyngien Pharyngeal endoblast
Enzymes Enzymes
Epiblaste Epiblast
Espace périvitellin Perivitellin space
Face dorsale Dorsal face
Face ventrale Ventral face
Fibres longitudinales Longitudinal fibers
Formation de l’axonème du flagelle (D) Formation of flagellar axoneme (D)
Formation des vésicules proacrosomiques (B) Formation of proacrosomalvesicles (B)
Gaine fibreuse Fibrous sheath

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Gangue gélatineuse ou muqueuse Gelatinous or mucous gangue
Globules polaires Polar globules
Gradient vitellin Vitelline gradient
Granule cortical Cortical granule
Granules pro-acrosomiques Pro-acrosomal granules
Lumière du tube séminifère Lumen of the seminiferous tubule
Manchon mitochondrial Mitochondrial sheath
Matériel cordal Cordal material
Matériel préchordal Prechordal material
Membrane basale Basement membrane
Membrane coquillère Shell membrane
Membrane de fécondation Fertilization membrane
Membrane ovulaire Ovular membrane
Membrane plasmique ou membrane ovulaire Plasma membrane or ovular membrane
Membrane vitelline Vitelline membrane
Mésoblaste Mesoblast
Mésoblaste dorsal Dorsal mesoblast
Mésoblaste ventral Ventral mesoblast
Microvillosités Microvilli
Microvillosités ovocytaires Oocyte microvilli
Mitochondrie Mitochondrion
Neurectoblaste Neurectoblast
Noyau Nucleus
Noyau Nucleus
Noyau de fécondation Fertilization nucleus
Noyau de l’ovule Nucleus of the ovum
Noyau en cours de condensation Condensing nucleus
Œuf alécithe Alecite egg
Œuf mégalécithe et centrolécithe Megalocith and centrolocith egg
Œuf mégalécithe et télolécithe Megalocith and telolecith egg
Œuf mésolécithe Mesolecith egg
Œuf oligolécithe Oligolecite egg
Ovocyte I (2n) Oocyte I (2n)
Ovocyte II (n) Oocyte II (n)
Ovocyte II en métaphase II Oocyte II in metaphase II
Ovotide ou ovule (n) Ovoid or ovule (n)
Phase d’accroissement Growth phase
Phase de différenciation (spermiogénèse) Differentiation phase (=spermiogenesis)
Phase de maturation (méiose) Maturation phase (meiosis)
Phase de multiplication (mitoses) Multiplication phase (mitoses)
Pièce intermédiaire Middle piece
Pièce terminale End piece
Plaque basale Basal lamina
Plaquette vitelline Vitelline plate
Pôle animal Animal pole
Pôle végétatif Vegetative pole
Pore de l’acrosome Acrosome pore
Protéines de structure Structural proteins
Queue Tail
Spermatide Spermatid
Spermatocyte I Spermatocyte I
Spermatocyte II Spermatocyte II
Spermatogonie Spermatogonia

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Spermatogonie croûtelleuse Crusty spermatogonia
Spermatogonie poussiéreuse pâle Pale dusty spermatogonia
Spermatogonie poussiéreuse sombre Dark dusty spermatogonia
Spermatogonie souche Sperùatogonia
Spermatozoïdes Spermatozoon
Tête Head
Tête spermatique Spermatic head
Traînée spermatique Sperm trail
Vaisseau sanguin Blood vessel
Vésicule acrosomique acrosomal vesicle
Vésicule golgienne Golgi vesicle
Vitellus Vitellus
Zone pellucide Zona pellucida

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