Lecture Notes

(Developmental Biology)

I. INTRODUCTION

A. Historical Background

Table 1. Some Scientists and Their Contribution to the Development of Embryology into a Science
Scientist Contribution
Aristotle - Secured information on embryos some of which are accurately
correct; his work symbolizes the beginning of the turning o f the
human mind away from superstition and conjecture and toward
observation;
- Attempted to give solution to the problem of ontogeny along the
general lines of his philosophical teachings, distinguishing between
the “substance” and the “form” of things; in turn, implied the
necessity of fertilization
Galen Learned much about the structure of advanced fetuses
de Graaf First to describe the ovarian follicle
Anton van Leeuwenhoek Invented the microscope
Hamm and Leeuwenhoek First saw the human sperm
Bonnet Discovered that eggs of some insects can develop
parthenogenetically
Spallanzani His work served as the initial step in bringing the experimental
method to bear on embryological problems; through a series of
ingeniously planned experiments he demonstrated that both the
female and the male sex products are necessary for the initiation
of development
Wolff Contended that embryological development occurs through
progressive growth and differentiation
Baer Also demonstrated the existence of the germ layers in embryos
Schleiden and Schwann First to contend the fact that the adult body is composed entirely of
cells and cell products; this paved the way for the realization of the
basic fact of embryology, “the body of the new individual is
developed from a single cell that is formed by the union, in
fertilization, of a germ cell contributed by the male parent with a
germ cell contributed by the female parent”

Thus, embryology began to become a science only until the introduction of the observation, invention of the
microscope, the advent of the experimental method, and the discovery of the cellular structure of the body.

The Prevailing Theories on Ontogentic Development of that time

1. Theory of Preformation
- claims that if one sees something developing from the egg, then that something must actually have been
there all the time but in the invisible form
ex. 1. the leaves, and sometimes the flower with all their parts can be discovered long before the bud
starts growing
2. the parts of the butterfly’s body (legs, wings, etc.) can be discovered if the cuticular coat of the
chrysalis is carefully removed a few days before the butterfly emerges

- all the parts of the embryo were imagined to be already present in the egg, but were thought to be
transparent, folded together and very small, so they could not be seen
- when the embryo began to develop, these parts supposedly started to grow, unfold and stretch themselves
becoming denser and therefore, visible
- the embryo, and indirectly the future animal, is preformed in the egg

2. Theory of Epigenesis by Wolff

- states that in the early egg, there does not exist a preformed embryo but only the materials of which the
embryo is built; the material does not represent an embryo
- the substance of which the embryo is formed is granular (presumably the cells or nuclei); the granules were
later arranged into layers (germinal layers); the body of the embryo is formed from the transformation of the
layers through the formation of local thickenings in some parts of these layers, by the thinning out in others,
and by the formation of folds and pockets
- there had to be an architect who would use the materials; the architect in a developing embryo is represented
by a “vital force”, perhaps not essentially different from the “creative principle” of Aristotle
B. Phases of Ontogenetic Development (An Overview)

1. Gametogenesis
- production of sex cells (gametes) in sexually-producing organisms through meiosis
a. spermatogenesis – production of spermatozoa in male sexually-producing organisms
b. oogenesis – production of ova in female sexually-producing organisms
- creates the conditions from which the subsequent embryogenesis takes its start
- in both sexes, the initials cells giving rise to the gametes are very similar and, as a rule, are not essentially
different from other cells of the body except that these cells are not involved in any of the differentiations
serving to support the life of the parent individual
- the first step in the production of gametes is a more or less rapid proliferation of cells by ordinary mitosis
(spermatogonia in the testes and oogonia in the ovaries); once proliferation ceases, the cells are called
spermatocytes and oocytes respectively; these then enter into a stage of growth and later into a stage of
maturation both stages differ to a very great extent in the two sexes
- the cytoplasmic differentiations of the spermatozoon enable it to reach the egg by active movement and to
fertilize it; the egg cell, however, accumulates in its cytoplasm substances which are used up during
development --- either directly by being transformed into the various structures of which the embryo consists,
or indirectly, as sources of energy for development; the elaboration in the egg cell of cytoplasmic substances
to be used by the embryo and their placing in correct positions are essential parts of what occurs during the
first phase of development
- meiosis discards half of the chromosomes, thus, singles out the set of genes that operate in the development
of a particular individual

2. Fertilization
- involves independent biological and physiological processes:
a. first, the spermatozoa must be brought into proximity with the eggs if fertilization is to occur; this
involves adaptations on the part of the parents which insure that they meet during the breeding
season
b. next, the spermatozoa must find the egg and fuse with it; this sperm-egg fusion entails a very
finely adjusted mechanism of morphological and physiochemical reactions
- the egg is then activated by a spermatozoon and starts developing
- among the first changes that take place in the egg after fertilization is the further rearrangement of the organ-
forming substances in the egg

3. Cleavage
- the fertilized egg undergoes a series of mitotic cell divisions in rapid succession
- during this period, the cleavage cells or blastomeres become smaller and smaller with each division
increasing the number of cells, but the size of the embryo does not change
- no far-reaching changes in the substances of the developing embryos can be discovered during this stage, as
if the preoccupation of the embryo seems to be in increasing the cell numbers
- the whole cleavage process is dominated by the cytoplasmic organoids of the cells, the centrosomes and
achromatic figures
- the nuclei multiply but do not interfere with the processes going on in the cytoplasm
- the result is sometimes a 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 blastocoele

4. Gastrulation
- during this phase, the single layer of cells, the blastoderm, gives rise to two or more layers of cells known as
the germinal layers; these are complex rudiments from which are derived the various organs of the animal’s
body
 in higher animals, the body consists of several layers of tissues and organs (e.g., the skin, etc.)
all of which may be traced to the 3 primary germinal layers
- of the three germ layers, the outermost, the ectoderm, always gives rise to the skin epidermis and the
nervous system
- the next layer, the mesoderm, is the source of the muscles, the blood vascular system, the lining of the
secondary body cavity (the coelom where such occur), and the sex organs
 in many animals, particularly the vertebrates, the excretory system and most of the internal
skeleton are derived from the mesoderm
- the third and innermost layer, the endoderm, forms the alimentary canal and the digestive glands
- the germinal layers are produced by the disappearance of a part of the blastoderm from the surface and its
enclosure by the remainder of the blastoderm
 the part that remains on the surface becomes ectoderm; the part disappearing into the interior
becomes endoderm and mesoderm
 the disappearance of the endoderm and mesoderm from the surface sometimes takes the form
of a folding-in of part of the blastoderm, so that the simple spherical body becomes converted into
a double-walled cup

5. Organogenesis
- this is the process of subdivision of the embryo into parts with specific destinies
- the continuous masses of cells of the 3 germinal layers become split up into smaller groups of cells, each of
which is destined to produce a certain organ or part of the animal
 every organ begins its development as a group of cells segregated from the other cells of the
embryo, the group being called the rudiment of the respective organ
  the rudiments into which the germinal layers become subdivided are called primary organ
rudiments some of which are very complex, containing cells destined to produce a whole system
of organs, such as the entire nervous system or the alimentary canal
 these complex primary organ rudiments later become subdivided into secondary organ
rudiments
- with the appearance of primary and secondary organ rudiments, the embryo begins to show some similarity to
the adult animal, or to the larva if the development includes a larval stage

6. Growth and Histological Differentiation

- the organ rudiments begin to grow and greatly increase their volume; in this way the animal gradually
achieves the size of its parents
- sooner or later the cells in each rudiment become histologically differentiated; that is, they acquire the
structure and physicochemical properties which enable them to perform their physiological functions
- when the cells in all the organs, or at least in the vitally important organs, have become capable of performing
their physiological functions, the young animal can embark upon an independent existence --- an existence in
which it has to procure food from the surrounding environment
 in rare cases (in nematodes, for instance) the young animal emerging from the egg is a
miniature copy of the adult animal and differs from the latter only in size and the degree of
differentiation of the sex organs; in this case the subsequent development consists only of growth
and maturation of the gonads
 in other cases, the animals emerging from eggs differ from the adult to a greater extent (i.e., not
only do the gonads but also the other organs may not be fully differentiated, or they even be
absent altogether and have to develop later; sometimes the animal emerging from the egg
possesses special organs which are absent in the adult but which are necessary for the special
mode of existence of the young animal); in this case the young animal is called a larva
 the larva undergoes metamorphosis when it is transformed into an animal similar to the adult;
the metamorphosis involves more or less drastic changes in the organization of the larva,
depending on the degree of difference between the larva and the adult; during metamorphosis
new organs may develop, so that the morphogenetic processes become active again after a more
or less prolonged [period of larval life
- a secondary activation of morphogenetic processes may be produced in a different way: many animals
possess considerable plasticity and may be able to repair injuries sustained from the environment or caused
experimentally
 lost parts may be regenerated, and this means that the 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 the adult stage

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