DEVELOPMENTAL BIOLOGY

Definition-Is the study of the process by which organisms grow and develop

 Modern developmental biology studies the genetic control of cell growth, differentiation
and “Morphogenesis” process that give rise to tissues, organs and anatomy.

One of the amazing conclusions of modern biological research is that the mechanisms of
development are very similar for all animals, including humans. This became possible due to
study of molecular basis of developmental processes. This has made developmental biology to be
the most exciting areas of biological research.

These dramatic advances came from three main traditions that became fused together into a
single world view; experimental embryology, developmental genetics and molecular biology.

Experimental Embryology;

Has been in existence since the beginning of the 20 th Century, consisting mainly of micro-
surgical experiments on embryos of frogs and sea urchins.These demonstrated the existence of
embryonic induction: chemical signals that controlled the pathways of development of regions of
cells within embryos.

Developmental genetics:

Has existed for a long time but thrived in the late 1970s when mass genetic screens were carried
out on fruit fly, Drosophila, in which thousands of mutations affecting development were
examined. These mutagenesis screens resulted in identification of a high propotion of the genes
that control development, not just in Drosophila, but in all animals.

Molecular Biology:

Started with the discovery of the three dimensional structure of DNA in 1953, and became a
practical science of gene manipulation in the 1970s. The key technical innovations were methods
for molecular cloning to enable single genes to be amplified to a chemically useful quantity,
methods for nucleic acid hybridization to enable the identification of DNA or RNA samples and
methods for DNA sequencing to determine the primary structure of genes and their protein
products. Once this tool kit had been assembled it could be applied to a whole range of biological
problems, including those of development. It was initially used to clone the developmental genes
of Drosophila. Most of key Drosophila genes were found to exist also in other animals and
frequently to be controlling similar developmental processes. Molecular biology methods were
also applied directly to vertebrate embryos and used to identify the previously mysterious
inducing factors and the genes regulated by them.

The application of molecular biology meant that mechanisms of development could be worked
out in molecular detail. It also meant that the path of development could be experimentally
altered by the introduction of new genes, or selective removal of genes,or by an alteration of the
regulatory relationships between genes. It has also been found that all animals use very similar
mechanisms to control their development. This means that one can learn about human
development by understanding how it happens in fruitfly, zebrafish, frog or mouse.

Position of Developmental Biology in Biology

Developmental biology occupies a pivotal position in modern biology. It unites the disciplines
of molecular biology, cell biology, genetics and morphology.

Molecular and cell biology is about how individual components work; the inducing factors, their
receptors, the signal transduction pathways, transcription factors. Genetics tells us directly about
the function of an individual gene and how it relates to activities of other genes.

Morphology or anatomical structure is both a consequence and a cause of the molecular events.
Hence developmental biology is a synthetic discipline in which an understanding of molecular
biology, genetics, and morphology is necessary.

Perspectives/ impact on society

 Development of a new life represents a new masterpiece of temporal and spiral control of
gene expression.
 Developmental genetics studies the effects that genes have in phenotype, given normal
or abnormal epigenetic parameters
 Findings of developmental biology has led to understanding that human embryos are
sensitive to damage during period of organogenesis and hence chromosomal basis of
some human birth defects like chromosomal aberrations that cause Down syndrome.
This can be detected in cells taken from amniotic fluid and form the basis of
amniocentesis tests taken by millions of expectant mothers every year. Many more
defects are due to mutations in genes that control development.
 An understanding of cell specialization during embryogenesis has provided information
on how stem cells specialize into specific tissues and organs- This information has led to
cloning of specific organs for medical purposes
 In vitro fertilization (IVF) is now a routine procedure and has enabled many previously
infertile couples to have a baby; its variants include; artificial insemination by donor
(AID),egg donation, and storage of fertilized eggs by freezing. AID,IVF, embryo freezing
and embryo transfer between mothers is also very important for farm animals. It has been
used for many years in cattle to increase the reproductive potential of the best animals.
 Another biological important process that occur during development is apoptosis-
Programmed cell death/suicide
 Many developmental models are used to elucidate the physiology and molecular basis of
the cellular process
  A deeper understanding of developmental biology can fasten the greater progress in
treatment of congenital disorders and diseases
 studying human sex determination can lead to treatment for disorders such as congenital
adrenal hyperplasia

Developmental Model Organisms

 Model Organisms used in developmental biology include:

 Frog- Xenopus leavis, Xenopus tropicalis
 Chicken- Caallus gallus
 Mouse- Mus musculus (mammalian embroyogenesis)
 Invertebrates
-Sea urchin
-Roundworm
-Fruitfy- Drosphila melanogaster(Drosophila embryogenesis)
 Plants(Plant embryogenesis)
- Vegetables
-Maize
 Others
-Slime Mold
Studied Phenomena:
Cell differentiation
 Formation of cell types from originally one cell- Zygote/Spore

Embryonic development

 Embryogenesis- is the steps in life cycle after fertilization- development of the

embryo, starting from zygote(fertilized egg)

Growth

 Is the enlargement of a tissue or organism.

 Growth continues after embryonal stage, and occurs through cell proliferation-
enlargement of cells or accumulation of extracellular material

Metamorphosis

 Most animals have larval stage with a body plan different from that of adult organism
 The larva develops into adult in a process called Metarmorphosis
Regeneration
 Is the activation of development so that a wigging body part grows back
 The phenomenon has been studied in salamanders, where the adults can
reconstruct a whole limb after it has been amputated
  Researchers hope to one day be able to induce regeneration in humans(i.e in
regenerative medicine)
 There’s little spontaneous regeneration in adult humans through liver i.e an
exception
 The regeneration of the liver involves dedifferentiation of some cells to a more
embryonal stage

ONTOGENIC DEVELOPMENT AS SUBJECT MATTER OF EMBRYOLOGY

Embryology-Study of ontogenic development of organisms

 In sexual reproduction new individual are produced by special generative cells or

gametes
 Gametes are different from other cells of animals because they go through the process of
maturation or meiosis as a result of which they lose ½ of their chromosomes and become
haploid, whereas other cells i.e. somatic cells are diploid.
 In multicellular animals, exist two types of sex cell: female; Ova and male; spermatozoa
 They unite in process of fertilization before development can start
 They fuse to form zygote with diploid number of chromosomes
 Zygote develop into new adult
 In asexual reproduction offspring are not derived from generative cells (gametes) but
from parts of parent’s body consisting of somatic cells.
 In sexual reproduction offspring develop from egg
 Hence to distinguish two forms of ontogenic development, the term “embryogenesis”
may be used to refer to develop from egg and Blastogenesis may be used for
development of individual through asexual reproduction

PHASES OF ONTOGENIC DEVELOPMENT

 Process leading to the development of a new individual starts before fertilization of egg,
because ripening of the egg and formation of the spermatozoa constitutes phase of
gametogenesis, create conditions for which subsequent embryogenesis starts
 In both oogenesis and spermatogenesis, meiosis by discarding ½ of chromosomes
singles out, set of genes involved in development of a particular individual
 In both sexes the initial cells that give rise to gametes are similar, as a rule, not different
from other cells of the body except they are not involved in any of the differentiations
serving to support life of the parental individual
 In both sexes 1st step in gamete formation is rapid proliferation of cells by ordinary
mitosis
 In male these proliferating cells are known as spermatogonia and in female, Oocytes
 They then enter stage of growth then maturation
 The process of growth and maturation however differs in two sexes i.e. cytoplasmic
differentiation of spermatozoa enable it reach egg by active movement and fertilize it
 Egg cell accumulate in its cytoplasm substances used up during development either
directly by becoming transformed into various structures of which embryo consists, or
indirectly as a source of energy for development
 Elaboration in the egg cell of cytoplasmic substances to be used by embryo and their
placing in correct positions are essential parts of what occurs during 1st phase of
development
 2nd phase of development is fertilization
 It involves a number of independent biological and physiological processes
 1st the spermatozoa must be brought closer to the egg if fertilization is to occur; this
involves adaptations on part of the parents which ensures that they meet during
breeding season, discharge sex cells simultaneously in case of external fertilization or
copulate incase of internal fertilization
 2nd spermatozoa must find the egg and fuse with it
 3rd phase of development is period of cleavage
 The fertilized egg is still a single cell, since nucleus and cytoplasm of spermatozoon
fuse with nucleus and cytoplasm of the egg
 If a complex multicellular organism is to develop from single cell, the egg, the latter
must give rise to a large number of cells by a number of mitotic cell divisions in
quick succession
 During this period the size of the embryo does not change, the cleavage cells or
blastomere become smaller and smaller with each division
 The whole process of cleavage is dominated by the cytoplasmic organoids of the the
cells, the centrosomes and a chromatic figures
 The nuclei multiply but do not interfere with the processes going on in cytoplasm
 Results of cleavage is at times compact heap of cells, but usually the cells are
arranged in a hollow spherical body, a blastula, with a layer of cells, the blastoderm,
surrounding a cavity the blastocoels
 (Fig)
 4th phase is Gastrulation
 During this phase, single layer of cells, the blastoderm gives rise to two or more
layers of cells known as Germinal layers
 Germinal layers are complex rudiments from which various organs of animal body
are derived
 In higher animals the body consists of several layer of tissues and organs e.g. skin,
subcutaneous connective tissue, layer of muscles, wall of gut e.t.c
 All these tissues and organs may be traced back to three layers (germinal layers) of
the cells
 Of these the external one, the ectoderm, always gives rise to skin epidermis and
nervous system
 Layer next to ectoderm, mesoderm is a source of muscles, the blood vascular system,
the lining of secondary body cavity (the coelom) in animals in which such a cavity is
present, the sex organs
 In vertebrates, the excretory system and most of the internal skeleton are also derived
from mesoderm
 The 3rd and innermost germinal layer, the endoderm, forms alimentary canal and
digestive glands
 The germinal layers are produced by the disappearance of a part of the blastoderm
from the surface and its enclosure by remainder of the blastoderm
 The part that remains on surface becomes ectoderm
 The part disappearing into the interior becomes endoderm and mesoderm
 Disappearance of endoderm and mesoderm from surface sometimes takes the form of
an in-folding of part of blastoderm, hence the spherical body becomes converted into
a double walled cup as if one side of the wall of an elastic hollow ball had been
pushed in by an external force
 The infolding of mesoderm and endoderm is known as invagination and the resulting
embryo is known as gastrula- hence term gastrulation
 If the gastrula is formed by invagination, the cavity of the double walled cup is called
the archenteron and the opening leading from this cavity to the exterior is called
blastopore
 In animals where gastrula is not formed by invagination, the cavity(archenteron) and
opening of cavity to exterior(blastopore) may still appear later
 The archenteron or part of it finally gives rise to the cavity of the alimentary system
 Fate of blastopore differs in three main groups of metazoan
o In Coelenterates-it becomes oral opening
o In Protostomia(annelids, Molluses, arthropods and allied groups)-its
subdivided into two openings, one which becomes mouth and other anus
o In Deutrostomia(echinoderms and chordates)
-blastopore is connected to development of anal opening
 -the mouth is formed later on as an independent perforation of the body wall
-the whole body lining of the alimentary canal does not always consist of
endoderm in all groups of animals
-the ectoderm may be invaginated secondarily at the oral or at both oral and
anal openings to become a part of the alimentary canal
 The parts of the alimentary canal lined by ectoderm are known as the stomodeum
(adjoining the mouth) and proctodeum (adjoining anus)

 The 5th phase of development- is subdivision of embryo into parts with specific
destinies i.e. phase of organogenesis(organ formation)
 The continuous masses of cells of three germinal layers become split into smaller
groups of cells each destined to produce a certain organ or part of the animal
 Every organ begins its development as a group of cells segregated from other cells
of embryo, this group of cells is referred to as rudiment of respective organ
 The rudiments into which germinal layers become subdivided are called primary
organ rudiments some are complex containing cells destined to produce whole
system of organs i.e. entire nervous system or alimentary canal
 These complex primary organ rudiments later subdivided into secondary organ
rudiments- the rudiments of the subordinated and simpler organs and parts
 Process of formation of primary organ rudiments follows closely the process of
gastrulation, hence the two cannot be considered separately
 With appearance of primary and secondary organ rudiments, the embryo begin to
show some similarities to adult animal or larva if development includes larval
stage

 6th phase of development is period of growth and histological differentiation.

 After the organ rudiments are formed they begin to grow and increase in volume and
gradually achieves size of its parent
 Sooner or later the cells in each rudiment become histologically differentiated i.e.
acquire the structure and physicochemical properties which enables them to
perform physiological functions, hence young animals can acquire food from
surrounding environment
 In rare cases(in nematodes) the young emerging from egg is a miniature copy of
the adult and differs only in size and degree of differentiation of sex organs, hence
subsequent development consist only of growth and maturation of gonads
 Sometimes animals emerging from egg has special organs which are absent in
adult but are necessary for the special mode of existence of young animal
  In this case young animal is called larva, larva may lead a different mode of life
from adult and therein lies one of advantages of having larval stage in
development
 The larva undergoes metamorphosis to transform into an animal similar to the
adult
 Morphogenetic processes become active again after a more or less prolonged
period of larval life
 A 2° activation of morphogenic processes may be produced in a different way
 Many animals posses considerable plasticity and may be able to repair injuries
sustained from environment or caused experimentally
 Lost parts may be regenerated hence developmental processes may sometimes be
repeated in an adult or adolescent organism
 Asexual reproduction of animals involves the development of new parts and
organs in animals that have already achieved adult stage

TRENDS OF THOUGHT IN EMBRYOLOGY

Baer’s Law-states that

“More general features that are common to all the members of a group of animals are in the
embryo, developed earlier than the more special features which distinguish the various members
of the group”

 Thus features that characterize all vertebrate animals(brain, spinal cord, axial skeleton in
form of notochord, segmented muscles, aortic arches) are developed earlier than the
features distinguishing the various classes of vertebrates(limbs in quadrupeds, hair in
mammals, feathers in birds etc)
 The characteristics distinguishing the families, genera and species come last in the
development of individual
 The early embryo thus has a structure common to all members of a larger group of the
animal kingdom and may be said to represent the basic plan of organization of that
particular group
 The group having a common basic plan of organization are the phyla of animal kingdom
 Baer’s law was formulated at a time when the theory of evolution was not recognized by
majority of biologists
 The law can now be reinterpreted in the light of evolutionary theory
 In its new form its known as Biogenetic law of Muller-Haeckel
 Muller-Haeckel propounded the law and supported it by extensive observations on
development of crustaceans(1864)
  According to Baer’s law common features of large groups of animals develop earliest
during ontogeny
In evolutionary theory, these features are ones that are inherited from common ancestors
of the animal group, in question thus they have an ancient origin
 Features that distinguish various animals from one another are those that they acquire
later in course of evolution
 Baer’s law states- that these features in ontogeny develop at later stages i.e. features of
ancient origin develop early in ontogeny; features of newer origin develop later
Hence ontogenetic development presents the various features of the animal’s
organization in the same sequence as they evolved during the phylogenetic development
“Ontogeny is a recapitulation of phylogeny”
-But the repetition is not a competitive one and biogenetic law states that
“Ontogeny is a shortened and modified recapitulation of phylogeny”

Why it’s a shortened process

1. What had once taken thousands of millions years(phylogeny) is new performed in days
and weeks(Ontogeny)
2. Many stages which occurred in original phylogenetic development may be omitted in
ontogeny

Why it’s a modified process

 The embryo at any given time is a living system which has to be in harmony with its
surroundings if it is to stay
 Embryo must be adapted to its surrounding and these adaptations necessitate the
modification of inherited features of organism e.g. the presence of placenta in mammals
*Placenta is a structure developed by embryo to establish a connection with uterine wall
of the mother to provide nutrition of embryo. This structure never existed in adult
mammalian ancestor
 The fragmentally repetition of certain ancestral characters may be useful in elucidating
the relationship of animals e.g.- formation of gill clefts or at least pharyngeal pouches in
ontogenic development of all vertebrates
 In aquatic vertebrate e.g. cyclostomata and fishes, the gill clefts serve as respiratory
organs
 In adult state of terrestrial vertebrate the pharyngeal pouches have disappeared
completely or have been modified out of all recognition, and the function of respiration
has been taken over by other organs e.g. lungs. But pharyngeal pouches appear in embryo
 In amphibians whose larvae are aquatic, the pharyngeal pouches serve temporarily for
respiration
  In reptiles, birds and mammals pharyngeal pouches of embryo do not serve for
respiration but its an indication that terrestrial vertebrate have evolved from aquatic forms
with functional gills

EXPLAINING DEVELOPMENT

Theories of Preformation and Epigenesis

Fundamental questions presented by existence of cyclical ontogenetic changes are:

1) Why does ontogeny occur at all?

2) What are the forces which produce changes?
3) How is it that starting from a simple spherical cell, the process always ends in producing
a highly complex and specific structure which though varying in detail, reproduces with
preservation of the same or almost same adult form?
 In the C17th and C18th when all biological science developed rapidly, together with the
physicochemical sciences, a theory was coined to explain the ontogenetic development of
animals i.e. theory of preformation

The Theory of Preformation

Claims that if we see that something develops from the egg, then this something must actually
have been there all the time but in an invisible form.

 All the parts of future embryo were imagined to be already in the egg, but they were
thought to be very transparent, folded together and very small, so that they could not be
seen when embryo began to develop these parts supposedly started to grow, unfold and
stretch themselves, become denser and thus more readily visible. The embryo and thus
indirectly the future animal was preformed in the egg. Hence theory of preformation
(reference Ovists and animal culist argument)
 The theory of preformation, though popular in its time, did not satisfy all biologists and
opposing views denying existence of a preformed embryo in the egg were proposed
 Hence came the theory of epigenesis by Caspar Friedrich Wolf(1759)
 Wolf adduced his own observations on the formation of chick embryo
 In earliest stages of development of the chick, he could not find any parts of future
embryo
 He found that egg was devoid of any visible structure there was a structure present but
different from that of later embryo
 He found that embryo is composed of granular substance presumably the granules could
have been cells or the nuclei
 These granules were later arranged into layers which we now call germinal layers
  Wolf saw that by the formation of local thickenings in some parts of these layers, by
thinning out in others and by the formation of folds and pockets, the layers are
transformed into body of the embryo
 He concluded that in the early egg, there does not exist a preformed embryo but only the
material of which embryo is built

Experimental Embryology

Allogamy – cross fertilization

Autogamy – self fertilization

Parthenocarpy – development of embryo without fertilization