CHAPTER 1 ● Published De Generatione Animalium in
Early Philosophers: Ancient Greek philosophers 1651, promoting the idea that "all things
like Democritus and Pythagoras theorized on sex come from the egg" (Ex ovo omnia).
determination (e.g., males from right testicle, ● His work was influenced by Aristotle's view
females from left). that the male semen provided a formative
Aristotle (384–322 BC): force to menstrual blood.
● Mistakenly concluded that no embryos were
● Classified animals by birth type: oviparity present in the uterus for several days
(egg-laying), viviparity (live birth), and post-coitus based on his studies in deer.
ovoviviparity (eggs hatch inside the body). ● First to describe the blastoderm in chick
● Described holoblastic and meroblastic cell embryos and early stages of blood formation.
division patterns.
● Concept of epigenesis: organ systems Marcello Malpighi (1628–1694):
develop gradually, not preformed.
● Published the first microscopic account of
Ernst Haeckel (1834–1919): chick development in 1672.
● Observed the neural groove, somites, and
● Proposed "Ontogeny recapitulates blood circulation in chick embryos.
phylogeny" (now debunked): development ● Supported preformation theory, believing a
mimics species evolution. preformed version of the chicken was in the
● Introduced the hypothetical ancestor egg.
"gastraea" to explain common stages like
the gastrula in metazoans. Regnier de Graaf (1641–1673):
Medieval Era: Kopho (13th century) used pigs for ● Performed detailed studies of female
anatomical studies due to religious restrictions on reproductive organs and considered
human dissection. mammalian ovarian follicles to be eggs.
Renaissance: ● His work later led to the naming of the
Graafian follicle.
● Leonardo da Vinci (1452–1519): Detailed ● Died young, delaying the discovery of the
drawings of the uterus and fetal membranes mammalian egg by 150 years.
(though inaccurately depicted human
placentas as ruminant-like). Karl Ernst von Baer (1792–1876):
● Invention of the printing press by Johann
Gutenberg enabled the spread of ● First to observe the mammalian egg in 1827,
anatomical knowledge. opening the "Graafian follicle" and examining
● Hieronymus Fabricius published De it under a microscope.
Formato Foetu in 1600, one of the first works ● Identified the small yellow spot as the
on comparative embryology. mammalian egg, advancing embryology.
Advances in the 16th Century: Bartolomeo Anton van Leeuwenhoek (1632–1723):
Eustachius (1514–1574) produced early
● Developed a microscope with 300x
illustrations of embryos, contributing to the field of
magnification and first observed moving
embryology.
spermatozoa in 1677.
Zacharias Janssen (1590): Invented the
● Described sperm as "animalcules" with thin,
microscope, which led to new discoveries in
undulating tails.
embryology.
● His findings led to debates, with some
William Harvey (1578–1657):
scientists (spermists) believing the sperm
head contained preformed miniature
organisms.
● Discovery of mammalian gametes sparked a in frogs, leading to the concept of genomic
debate on whether the embryo originates from totipotency.
the egg or sperm. ● John Gurdon later proved that somatic cells
● Two schools of thought emerged: epigenesis retain totipotency by performing serial nuclear
(embryo develops progressively) and transfers, although producing adult organisms
preformation (embryo is prefigured in the from somatic cells remained challenging in frogs.
gametes). ● Steen Malte Willadsen (1986): mammalian
● Preformation had religious and philosophical cloning, sheep, morula-stage, enucleated eggs,
support, as it negated the need for a "mysterious electrical fusion.
force" to initiate development. ● Keith H Campbell, Ian Wilmut (1996): Roslin
● Preformation also aligned with the idea that future Institute, lamb cloning, cultured cells, cell cycle
generations were preformed, similar to Russian synchronization.
nesting dolls, but lacked a size scale since cell ● Transgenic animals, Polly: human clotting factor
theory was not yet proposed. IX, biopharming.
● Caspar Friedrich Wolff (1734–1794) challenged ● Dolly (1997): mammary gland cells, 6-year-old
preformation through detailed observations on ewe, global attention.
chick embryos, supporting epigenesis. ● Quiescence: donor cells, G0 phase, later
● Preformation persisted until the 1820s when research, non-essential.
advances in microscopy and tissue staining ● Species cloning: cattle, mice, goats, pigs, dogs.
shifted focus to new embryological discoveries. ● Embryology: rapid evolution, genetics, veterinary
● Christian Pander (1794–1865) discovered germ education, 19th century.
layers (ectoderm, mesoderm, and endoderm) ● Key texts: Patten "Embryology of the Pig",
while studying chick embryos. Gilbert "Developmental Biology."
● Martin Heinrich Rathke (1793–1860) studied ● Terminology: Latin, Greek, modernized.
comparative embryology and identified
pharyngeal arches common to all vertebrates. CHAPTER 2
● Karl Ernst von Baer identified the mammalian ● Embryonic development begins with fertilization,
ovum and described the notochord, recognizing forming the zygote.
common developmental principles across ● Development continues from embryo to fetus,
species. involving organ formation and refinement.
● By the late 19th century, August Weismann ● The embryonic period involves major organ system
proposed that both egg and sperm contribute formation; the fetal period focuses on growth and
equally to inheritance via chromosomes. maturation.
● Theodor Boveri and others identified ● Development continues post-birth, with organs
growing and maturing until puberty.
chromosomes as distinct and responsible for
● Fertilization initiates with sperm penetration of the
different vital processes, linking them to sex
oocyte, forming the zygote with maternal and
determination. paternal pronuclei.
● In the early 20th century, genetics (studying ● Cleavage divisions increase cell number without
inheritance) and embryology (studying increasing overall embryo volume, forming smaller
development) diverged, leading to tensions blastomeres.
between the two fields. ● Morula forms from blastomeres; compaction leads to
● Salome Gluecksohn-Schoenheimer and trophectoderm formation.
Conrad Waddington advocated for a synergy ● Blastocyst forms with a fluid-filled cavity and inner
between genetics and embryology, studying how cell mass (ICM), which will form the embryo.
● Gastrulation creates a trilaminar embryonic disc with
mutations affected development.
ectoderm, mesoderm, and endoderm germ layers.
● Nuclear transfer experiments by Robert Briggs
● Amniotic folds form around the embryonic disc,
and Thomas King in the 1950s demonstrated leading to the amniotic cavity.
that somatic cell nuclei could direct development ● The embryonic period ends when most organ
systems are outlined.
● Differentiation: Specialized cell types form from less ○ DNA Methylation: Condenses
specialized cells through differential gene chromatin, silencing genes.
expression. ○ Histone Modifications: Acetylation
● Cell commitment has two phases: specification opens chromatin for transcription.
(reversible) and determination (irreversible). ○ Polycomb-Trithorax Regulation:
● Induction: Cell-to-cell signaling drives differentiation, Controls chromatin states to regulate
requiring cell competence during a critical period. gene activity.
● Hans Spemann showed optic vesicle interaction is ● Transcription Factors: Activate expression of
necessary for lens formation in surface ectoderm. genes crucial for development.
● Induction examples: optic vesicle for lens formation
and notochord signals (e.g., Sonic hedgehog) for Patterning
neuroectoderm development.
● Patterning: Organization of cells into tissues and
Signaling Molecules and Families organs.
● Body Axes:
● Paracrine Factors/Morphogens: Signaling ○ Cranio-Caudal Axis: Established during
molecules acting over short distances. gastrulation.
● Major Families: ○ Dorso-Ventral Axis: Present in
○ Fibroblast Growth Factor (FGF) blastocyst, further refined by signaling.
Family: Includes >20 related proteins. ○ Proximo-Distal Axis: Developed during
○ Hedgehog Family: Includes Sonic limb formation.
hedgehog (Shh), Desert hedgehog (Dhh), ● Morphogens: Signaling molecules that create
and Indian hedgehog (Ihh). gradients to guide cell differentiation and tissue
○ Wingless (Wnt) Family: At least 15 formation (e.g., Sonic hedgehog in neural tube
members, interact with Frizzled patterning).
receptors.
○ Transforming Growth Factor-β (TGF-β) Homeobox (Hox) Genes
Superfamily: Includes TGF-β, Activin,
Bone Morphogenetic Protein (BMP), ● Function: Control positional identity along the
Nodal, Glial-Derived Neurotrophic Factor cranio-caudal axis in development.
(GDNF), Inhibin, and Müllerian Inhibitory ● Fruit Fly (Drosophila melanogaster):
Substance (MIS). ○ Homeotic genes direct segmental
development; mutations (e.g.,
Cellular and Genomic Potency Antennapedia) can cause body parts to
develop in incorrect locations.
● Cellular Totipotency: Ability of zygote and early ● Mammals:
blastomeres to form all embryonic and ○ Hox genes are grouped into four clusters:
extra-embryonic tissues. Hox-A, Hox-B, Hox-C, Hox-D.
● Pluripotency: Ability to form all embryonic ○ Genes in the 3′ end of each cluster are
tissues but not extra-embryonic tissues; seen in expressed first, controlling anterior body
ICM and epiblast. parts; genes in the 5′ end control
● Genomic Totipotency: Ability of differentiated posterior parts.
cells (e.g., mammary gland cell) to contribute to ○ Hox genes are expressed in an
all tissues with the aid of reprogramming. overlapping pattern, providing a specific
● Multipotency and Unipotency: Germ layers ‘Hox code’ for positional identity.
(ectoderm, mesoderm, endoderm) become ○ Retinoic Acid: Gradient likely involved in
multipotent and then unipotent, forming specific Hox code induction during gastrulation.
tissue types. ○ Organizer Regions: Early-gastrula,
mid-gastrula, and late-gastrula regions
Molecular Control of Differentiation (including the primitive node) create
anterior-posterior gradients influencing
● Epigenetic Changes: Hox gene expression.
Morphogenesis CHAPTER 3
● Definition: Mechanism by which tissues and Female Reproductive System
organs are shaped.
● Examples: ● Ovaries: Produce oocytes; site of follicle
○ Neural Tube Formation: Involves cell development.
shape changes and proliferation, forming ● Tubular Genital Tract:
the neural tube from a flat neural plate. ○ Oviduct: Has three parts – infundibulum
○ Gastrulation: Formation of somatic germ (receives oocytes), ampulla, and isthmus
layers and germ cell line through cell (site of fertilization).
involution. ○ Uterus: Bicornuate (two uterine horns);
○ Programmed Cell Death (Apoptosis): has a short body in most species but a
Creates gaps between digits in hand- and long body in the mare. Contains cervix
foot-plates. with internal and external orifices.
○ Vagina and Vestibulum: Transition areas
Epigenetic Modifications and Life leading to the vulva.
Cycles
Puberty
● DNA Methylation:
● Onset: Marks the start of regular cyclic activity in
○ Primordial Germ Cells: Highly
the ovary and behavioral changes.
methylated; undergo demethylation as
● Before Puberty: Oocytes develop but are not yet
they enter the genital ridge.
ready for fertilization.
○ Gametogenesis: De novo methylation of
● After Puberty:
DNA; sex-specific methylation leads to
○ Hormonal Regulation: Signals from the
genomic imprinting.
brain (pineal gland, hypothalamus,
○ Post-Fertilization: Paternal genome
pituitary gland) trigger the production of
demethylated early, both genomes
fertilizable oocytes.
become equally demethylated by the
○ Gonadotropins: FSH and LH are released
morula and early blastocyst stages.
from the anterior pituitary, regulated by
Imprinted genes escape demethylation.
GnRH from the hypothalamus.
● X-Chromosome Inactivation:
○ Factors Influencing Onset: Age, body
○ Function: Dosage compensation for
weight, breed, nutrition, disease, season,
X-linked genes in females.
and proximity to a male.
○ Process: Initiated by the XIST gene,
leading to inactivation of one
Estrous Cycle
X-chromosome.
○ In ICM: Random inactivation of
● Phases:
X-chromosome; in trophectoderm,
○ Prooestrus: Follicle growth; estrogen
preferential repression of the maternal
levels rise.
XIST gene leading to paternal
○ Oestrus (Heat): Peak estrogen levels
X-chromosome inactivation.
trigger GnRH surge, stimulating LH
○ Parental Genome Conflict: Preference
release and leading to ovulation.
for paternal gene expression in
○ Metoestrus: Formation of corpus luteum;
extra-embryonic tissues, maternal gene
progesterone production increases.
expression in embryo-forming cells.
○ Dioestrus: Maximum progesterone
production; prepares endometrium for
potential pregnancy.
● Ovarian Follicle Transformation: Post-ovulation,
follicle cells form the corpus luteum, producing
progesterone.
○ In Pregnant Animals: Progesterone
maintains pregnancy.
○ In Non-Pregnant Animals: Absence of Hormonal Maintenance of Pregnancy and
pregnancy leads to luteolysis (corpus Reproductive Cyclicity
luteum regression) due to
prostaglandin-F2α, resuming the estrous Maternal Recognition of Pregnancy
cycle.
● Endometrial Prostaglandin-F2α: In pregnant
Unique Cases animals, the release of this hormone into the
bloodstream is blocked, leading to the
● Cat and Camel: GnRH surge leading to ovulation persistence of the corpus luteum. This process is
is induced by copulation. Cats may display crucial for maintaining pregnancy and involves
estrous behavior and accept males even after species-specific signals from the embryo that the
entering metoestrus. endometrium recognizes.
● Species-Specific Mechanisms:
○ Dog and Cat: The mechanisms for
corpus luteum regression in these
species are not well understood. In dogs,
the corpus luteum can remain functional
for up to two months. In cats,
progesterone production shifts to the
placenta later in pregnancy, making the
corpus luteum less critical.
Cyclicity Patterns in Domestic Animals
● Non-Seasonal Polycyclic Animals:
○ Pigs and Cattle: These animals
experience cyclic activity throughout the
year, only interrupted by pregnancy,
lactation, or pathological conditions.
● Seasonally Polycyclic Animals:
○ Mare: A ‘long-day’ breeder; peak cyclic
activity occurs from spring to autumn.
During winter, mares become
anoestrous.
○ Queen (Cat): Becomes anoestrous in
autumn and resumes cyclic activity with
increasing daylight.
○ Ewe and Doe (Small Ruminants):
‘Short-day’ breeders; exhibit cyclic
activity in autumn and early winter,
followed by a period of anoestrus.
● Monocyclic Animals:
○ Bitch (Dog): Experiences one or two
oestrous periods per year, separated by
long periods of anoestrus with no clear
seasonality.
Hormonal Sources of Progesterone During
Pregnancy
● Cattle:
○ Early Pregnancy: Corpus luteum is the
main source of progesterone.
○ Mid to Late Pregnancy: The placenta
takes over progesterone production from
around Day 120–150 up to Day 250.
● Horse:
○ Early Pregnancy: Several accessory
corpora lutea and the primary corpus
luteum produce progesterone until the
end of the third month.
○ Late Pregnancy: The placenta produces
progesterone from the end of the third
○
month until term.
● Ewe:
○ Early Pregnancy: Corpus luteum is the
major source of progesterone.
○ Late Pregnancy: The placenta becomes
the primary source of progesterone.
● Pigs, Goats, Dogs, and Cats:
○ Throughout Gestation: The corpus
luteum is the major source of
progesterone.
Prolactin and Pseudopregnancy
● Bitch (Dog):
○ Pseudopregnancy: Characterized by a
prolonged period of luteal function, with
overt or covert signs during metoestrus
and dioestrus. Prolactin is thought to play
a role in this phenomenon.
● Queen (Cat):
○ Corpus Luteum Persistence: Corpora
lutea persist in the absence of pregnancy,
but the queen returns to oestrus during
the season of cyclic activity.
