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Introductory lesson 1

29.11.2012

Origin
The multidisciplinary approach to the study of
development first arose before the turn of the
Twentieth Century as an integration of
Embryology
(Study of embryonic
development)

ytology
( Study of cellular
structure and function)

enetics
(Study of inheritance)

Definition

Developmental biology is the study of the process


by which multicellular organisms grow and develop
from its early immature forms (zygote, larva,
embryo , etc.) into an adult.

Modern developmental biology studies the genetic


control of cell growth, differentiation and
morphogenesis, which is the process that gives rise
to tissues, organs and anatomy (structure).

History (remember on the basis of


importance )

American leading cytologists Edmund Beecher Wilson (1882) at


Columbia University recognized that development of the embryo
is a demonstration of changes in individual cells.

Wilson recognized that the characteristics of an organism


gradually appear by utilization of the inherited information that is
located on the chromosomes.

In 1885, the German embryologist Wilhelm Roux removed a


portion of the medullary plate of an embryonic chicken and
maintained it in a warm saline solution for several days,
establishing the principle of tissue culture.

History

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Wilhelm Roux believed that the fertilized egg receives substances that
represent different characteristics of the organism, this "qualitative division"
fixes the fate of the cells

In 1888, Roux published the results of a series of defect experiments in


which he took 2 and 4 cell frog embryos and killed half of the cells of
each embryo with a hot needle. He reported that they grew into half-embryos
and the separate function of the two cells had already been determined. This
led him to propose his Mosaic" theory of epigenesis: after a few cell
divisions the embryo would be like a mosaic, each cell playing its own
unique part in the entire design.

Roux had manipulated embryos and observed the effects of these


manipulations on them. For this reason, many embryologists consider him to
be the "Father of Experimental Embryology."

Roux's experimental approach to


embryology

History

page 3

German embryologist, Hans Driesch (1892),


work with sea urchin embryos.
Instead of destroying one of the cells of the twocelled embryo, he separated the cells from oneanother and found that isolated cells at the fourcell stage also develop normally.
Thus, Driesch concluded that each cell retains
all the developmental potential of the zygote.

Sea urchin are small, spiny, globular animals

When Driesch separated the blastomeres from 4- and 8-cell embryos, each
isolated blastomere regulated its development so as to produce a complete
organism.

Manipulation of the Embryo

(A) an early amphibian embryo is split almost into two parts with a hair
loop.
(B) an amphibian embryo at a somewhat later stage receives a graft of a
small cluster of cells from another embryo at that stage.
Result: a single embryo to develop into a pair of conjoined (Siamese)
twins.

History
page 4
The Role of hereditary Material
in develpoment

Mendel's pea plant experiments conducted between 1856 and 1863


established many of the rules of heredity, now referred to as the laws
of Mendelian inheritance.
Although equal distribution of hereditary information to all cells had been
established in the late 1800's, its role in development remained a
mystery.
In1900, the significance of Gregor Mendel's work on heredity was finally
appreciated.
The other contribution was made by Theodor Boveri (1902) he
demonstrated that normal development is dependent upon the normal
combination of chromosomes.

From then the research on development


biology is going on..

Basic Anatomical Features

The similarities between animal species in the genes that


control development reflect the evolution of animals from a
common ancestor

The body must have been organized with a sheet of skin


covering the exterior, a mouth for feeding and a gut tube to
contain and process the food, muscles, nerves and other
tissues arranged in the space between the external sheet
of skin and the internal gut tube.

These features are common to almost all animals and they


correspond to basic anatomical scheme of development.

Blastula
After fertilization egg divides and produce a hallow sphere
of epithelial cells surrounding a cavity is known as blastula

A. Scanning Electron Micrograph showing a


blastula with initial epithelial cells inside

Gastrulation

The fertilized egg cell divides to form many smaller cells. These
cohere to create an epithelial sheet.
Much of this sheet remains external, constituting the ectoderm
the precursor of epidermis and nervous system.
A part of the sheet becomes tucked into the interior to form
endodermthe precursor of the gut and its appendages, such
as lung and liver.
Another group of cells move into the space between ectoderm
and endoderm, and form the mesodermthe precursor of
muscles, connective tissues, and various other components.
This transformation of a simple ball or hollow sphere of cells
into a structure with a gut is called gastrulation.

Sea urchin gastrulation

Fig.1: The formation of ectoderm, endoderm and mesoderm. ( B) A group of


cells loose from the epithelium to become mesoderm. (C) These cells crawl
over the inner face of the wall of the blastula (D) Epithelium is continuing to
tuck (push/insert) inward to become endoderm. (E-F) The invaginating
endoderm extends into a long gut tube. (G) The end of the gut tube makes
contact with the wall of the blastula at the site of the future mouth opening.

Role of Regulatory DNA in development

When animal species with similar body plansdifferent


vertebrates such as a fish, a bird and a mammal, is compared,
it is observed that corresponding genes usually have similar
sets of regulatory modules.
The same result is observed between different species of
nematode worm, or different species of insect.
The protein-coding sequences of vertebrate and invertebrate
are unmistakably similar, but the corresponding regulatory DNA
sequences appear very different.
Different body plans are produced mainly by changing the
program of the regulatory DNA.

Regulatory DNA defines the Program of


development

Observation of Cellular Interactions

To explain development in terms of cell behavior, it is


needed to be able to track the individual cells through all
their divisions, transformations, and migrations in the
embryo.

Cellular interaction can be observed by manipulating the


embryo

Cell lineage tracing in the early chick


embryo
(A,D) Two tiny dots of fluorescent dye, one
red, the other green, have been used to
stain small groups of cells in an embryo at
20 hours of incubation. The dots have
been placed on each side of a structure
called the node.
(B,E) Six hours later, some of the labeled
cells have remained at the node (which
has moved backwards), giving a bright
spot of fluorescence there, while other
cells have begun to move forwards
(C,F) After a further 8 hours, the body plan
is clearly visible, with a head at the front
end , a central axis, and rows of embryonic
body segments, called somites, on either
side.

Cell Makes Developmental


Decisions/Cell Fate

The ultimate differentiated state to which a cell has become


committed is called the cell fate.
The cell that is fated to become a neuron may be already
specialized in a way that guarantees that it will become a
neuron no matter how its surroundings are disturbed; such a
cell is said to be determined for its fate.
On the other hand the cell may be biochemically identical to
other cells destined for other fates, the only difference
between them being the accident of position, which exposes
the cells to different future influences.

Cell Fate

A cell may be already somewhat specialized for its normal


fate, but still able to change and undergo a different fate if it
is put in a sufficiently coercive environment.
Or the cell may be determined as a brain cell, but not yet
determined as to whether it is to be a neuronal or a glial
component of the brain. It seems, adjacent cells of the
same type interact and depend on mutual support to
maintain their specialized character.
Some developmental biologists would describe such a cell
is specified or committed, but not yet determined.

The standard test for cell fate


determination

Cells Have Remembered Positional


Values

In many systems, long before cells become committed


to differentiating as a specific cell type, they become
regionally determined: that is, they switch on and
maintain expression of genes that can be regarded as
markers of position or region in the body.

This position-specific character of a cell is called its


positional value, and it shows its effects in the way
the cell behaves in subsequent steps of pattern
formation.

Positional Values

In the chick embryo the leg and the wing originate at about the same
time in the form of small tongue-shaped buds projecting from the
flank (side).
The cells in the two pairs of limb buds appear similar and uniformly
undifferentiated at first.
But a simple experiment shows that this appearance of similarity is
unreliable.
A small block of undifferentiated tissue at the base of the leg bud,
from the region that would normally give rise to part of the thigh, can
be cut out and grafted into the tip of the wing bud.
Remarkably, the graft forms not the appropriate part of the wing tip,
nor a misplaced piece of thigh tissue, but a toe.

Positional Values

Therefore, the early leg-bud cells are already


determined as leg but are not yet permanently
committed to form a particular part of the leg:
they can still respond to changes in the wing
bud so that they form structures appropriate to
the tip of the limb rather than the base.

Normal condition

Morphogens Are Long-Range


Inducers in Cell Fate Determination

Signal molecules often seem to govern a simple


yesno choice: when their concentration is high or
low.

A signal molecule that imposes a pattern on a whole


field of cells is called a morphogen.

Morphogen mediated cell fate


determination

A group of cells at one side of the vertebrate embryonic


limb bud become specialized as a signaling center and
secrete Sonic hedgehog proteina member of the
Hedgehog family of signal molecules.
This protein spreads out from its source, forming a
morphogen gradient that controls the characters of the
cells along the thumb-to-little-finger axis of the limb bud.
If an additional group of signaling cells is grafted into
the opposite side of the bud, a mirror duplication of the
pattern of digits is produced

Sonic hedgehog as a morphogen in


chick limb development
(A) Expression of the Sonic
hedgehog gene in a 4-day chick
embryo. The gene is expressed
in the midline of the body and at
the posterior border of each of
the two wing buds. Sonic
hedgehog protein spreads out
from these sources.
(B) Normal wing development.
(C) A graft of tissue from the
polarizing region causes a
mirror-image duplication of the
pattern of the host wing.

Apoptosis Part of the Program of Development

Apoptosis, or programmed cell death (PCD), is a form of cell death


that is generally triggered by normal, healthy processes in the body
of multicellular organisms.
Biochemical events lead to characteristic cell changes (morphology)
such as release of proapoptotic proteins, shrinking of the cells
cytoplasm, nuclear fragmentation, chromatin condensation, increase
of mitochondrial membrane permeability, formation of cell fragments
called apoptotic bodies that phagocytic cells are able to engulf.
It is a series of programmed steps that cause a cell to die by self
digestion without releasing intracellular contents into the surrounding
environment.

Apoptosis

The control of cell numbers in development depends on cell death as well as cell
division.
If apoptosis is prevented in the body, cells will grow uncontrollably and cause
cancer.
For example, people with chronic myelogenous leukemia (CML) typically have
1025 times many white blood cells than normal.
Apoptosis confers advantages during an organism's lifecycle. For example, the
separation of fingers and toes in a developing human embryo occurs because
cells between the digits undergo apoptosis.
Caenorhabditis elegans generates 1030 somatic cell nuclei in the course of its
development, but 131 of the cells die by apotosis.
Thus programmed cell deaths occur in an absolutely predictable pattern.

Apoptosis

learn more from


Molecular Biology Of
The Cell - Alberts B. 5th Ed.
Chapter 18

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