BS Zoology 5th

ZOO-504 DEVELOPMENTAL BIOLOGY Cr. Hours: 4(3+1)

Introduction: Principal features of

development, origin of sexual
reproduction, developmental patterns;

Hafiz Mehmood Ul Hassan

Lecturer in Zoology, GPGC Nowshera
M.Phil Zoology/ Fisheries & Aquaculture (UoP)
M.Sc Zoology/Fisheries (UoP)
Genotoxicology (Humans)
M.Ed Science Education (AIOU)
[email protected]
0302-5772747
Principal features of development
 Developmental Biology
The discipline that studies embryonic and other developmental processes.
Development: It Is a relatively slow process of progressive changes.
In nearly all cases, the development of a multicellular organism begins with
a single cell the fertilized egg, or zygote, which divides mitotically to
produce all the cells of the body.
The study of animal development has traditionally been called embryology,
from that stage of an organism that exists between fertilization and birth.
But development does not stop at birth, or even at adulthood. Most
organisms never stop developing. Each day we replace more than a gram of
skin cells (the older cells being sloughed off as we move), and our bone
marrow sustains the development of millions of new red blood cells every
minute of our lives. In addition, some animals can regenerate severed parts,
and many species undergo metamorphosis (such as the transformation of a
tadpole into a frog, or a caterpillar into a butterfly). Therefore, in recent
years it has become customary to speak of developmental biology as the
discipline that studies embryonic and other developmental processes.
 Embryology
• The study of the developmental process from a
single cell to a baby in 9 months.
• In other words, investigations of the molecular,
cellular and structural factors contributing to
the formation of an organism.
 Principal features of development
• Development accomplishes two major objectives: it generates
cellular diversity and order within each generation, and it
ensures the continuity of life from one generation to the next.
Thus, there are two fundamental questions in developmental
biology:
– How does the fertilized egg give rise to the adult body, and
– How does that adult body produce yet another body?
• These two huge questions have been subdivided into six
general questions scrutinized by developmental biologists:
• Or they can be categorized as principles of development.
 Principal features. of development
• Differentiation. A single cell, the fertilized egg, gives rise to hundreds
of different cell types muscle cells, epidermal cells, neurons, lens cells,
lymphocytes, blood cells, fat cells, and so on. This generation of cellular
diversity is called differentiation. Since each cell of the body (with very
few exceptions) contains the same set of genes, we need to understand
how this same set of genetic instructions can produce different types of
cells. How can the fertilized egg generate so many different cell types?
• Morphogenesis. Our differentiated cells are not randomly distributed.
Rather, they are organized into intricate tissues and organs. These organs
are arranged in a given way: the fingers are always at the tips of our
hands, never in the middle; the eyes are always in our heads, not in our
toes or gut. This creation of ordered form is called morphogenesis. How
can the cells form such ordered structures?
 Principal features of development
• . to stop dividing? If each cell in
Growth. How do our cells know when
our face were to undergo just one more cell division, we would be
considered horribly malformed. If each cell in our arms underwent just
one more round of cell division, we could tie our shoelaces without
bending over. Our arms are generally the same size on both sides of the
body. How is cell division so tightly regulated?
– Such growth, in which the shape is preserved because all components grow at the
same rate, is called isometric growth. In many organisms, growth is not a uniform
phenomenon. It is obvious that there are some periods in an organism's life during
which growth is more rapid than in others. Physical growth during the first 10
years of person's existence is much more dramatic than in the 10 years following
one's graduation from college. Moreover, not all parts of the body grow at the
same rate. This phenomenon of the different growth rates of parts within the same
organism is called allometric growth (or allometry).
• Reproduction. The sperm and egg are very specialized cells. Only
they can transmit the instructions for making an organism from one
generation to the next. How are these cells set apart to form the next
generation, and what are the instructions in the nucleus and cytoplasm
that allow them to function this way?
 . of development
Principal features
• Evolution. Evolution involves inherited changes in development. When
we say that today's one-toed horse had a five-toed ancestor, we are
saying that changes in the development of cartilage and muscles
occurred over many generations in the embryos of the horse's ancestors.
How do changes in development create new body forms? Which
heritable changes are possible, given the constraints imposed by the
necessity of the organism to survive as it develops?
• Environmental integration. The development of many organisms is
influenced by cues from the environment. Certain butterflies, for
instance, inherit the ability to produce different wing colors based on the
temperature or the amount of daylight experienced by the caterpillar
before it undergoes metamorphosis. How is the development of an
organism integrated into the larger context of its habitat?
 Anatomical approaches to development.
• Comparative embryology, the study of how anatomy changes during
the development of different organisms. For instance, a comparative
embryologist may study which tissues form the nervous system in the
fly or in the frog.
• Evolutionary embryology, the study of how changes in development
may cause evolutionary changes and of how an organism's ancestry may
constrain the types of changes that are possible.
• Teratology, the study of birth defects. OR Study of the embryological
causes for birth defects (Genetic and Environmental). These anatomical
abnormalities may be caused by mutant genes or by substances in the
environment that interfere with development. The study of abnormalities
is often used to discover how normal development occurs.
Origin of Sexual Reproduction
 Origin of Sexual Reproduction
The evolution of sexual reproduction describes how sexually reproducing
animals, plants, fungi and protists could have evolved from a common
ancestor that was a single celled eukaryotic species. There are a few species
which have secondarily lost the ability to reproduce sexually, such as
Bdelloidea (Rotifers) , and some plants and animals that routinely reproduce
asexually (by apomixis and parthenogenesis) without entirely losing sex.

The origin of sexual reproduction in prokaryotes is around 2 billion years

ago when bacteria started exchanging genes via the processes of
conjugation, transformation, and transduction. In eukaryotes, it is thought to
have arisen in the Last Common Eukaryotic Ancestor (LECA), possibly via
several processes of varying success, and then to have persisted.
Why did sex evolve?

• Life originated without sex (as

best we can tell) so sexual
reproduction is something that
had to evolve

• There are a large number of

disadvantages to sexual
reproduction which makes the
evolution of sex a conundrum
 Sex is not necessary for all life
• Some plants and animals have entirely
abandoned sex

• Others have sex only when its convenient and

are asexual most of the time (facultatively
sexual)
Ancient asexuals: Bdelloid rotifers

• bdelloid rotifers date

back ~100 million
years

• Despite bdelloids'
asexuality, they've
diversified into 380
species
Facultative sexuality in animals
• In some animals, such as
Hydra, asexual reproduction
can occur through budding

• These animals are still capable

of reproducing sexually as well

• Sexual and asexual processes

are governed by environmental
conditions
 Parthenogenesis – offspring from
unfertilized eggs

Cnemidophorus velox, a parthenogenic lizard

Aphids – asexual and sexual

• Females give birth to live

females during the
summer months

• As winter approaches,
both males and females
are produced, which mate
to produce eggs
Evidences or Theories for origin of
Sexual Rep:

• Since hypotheses for the origins of sex are difficult to verify

experimentally (outside of evolutionary computation), most current
work has focused on the maintenance of sexual reproduction.
• The maintenance of sexual reproduction - specifically, of its
dioecious form - in a highly competitive world had long been one of
the major mysteries of biology, as both other known mechanisms of
reproduction - asexual reproduction and hermaphroditism - possess
apparent advantages over it.
• Many protists reproduce sexually, as do the multicellular plants,
animals, and fungi. In the eukaryotic fossil record, sexual
reproduction first appeared by 1.2 billion years ago in the
Proterozoic Eon.
• All sexually reproducing eukaryotic organisms likely derive from a
single-celled common ancestor. It is probable that the evolution of
sex was an integral part of the evolution of the first eukaryotic cell.
There are a few species which have secondarily lost this feature,
such as Bdelloidea and some parthenocarpic plants.
 Evidences or Theories : Diploidy
• Organisms need to replicate their genetic material in an efficient and
reliable manner. The necessity to repair genetic damage is one of the
leading theories explaining the origin of sexual reproduction. Diploid
individuals can repair a damaged section of their DNA via homologous
recombination, since there are two copies of the gene in the cell and if one
copy is damaged, the other copy is unlikely to be damaged at the same
site.
• A harmful mutation in a haploid individual, on the other hand, is more
likely to become fixed (i.e. permanent), since any DNA repair mechanism
would have no source to recover the original undamaged sequence from.
• The most primitive form of sex may have been one organism with
damaged DNA replicating an undamaged strand from a similar
organism in order to repair itself.
 Evidences or Theories : Meiosis
• If, as evidence indicates, sexual reproduction arose very early in eukaryotic
evolution, the essential features of meiosis may have already been present in
the prokaryotic ancestors of eukaryotes.
• Natural transformation in bacteria, DNA transfer in archaea, and meiosis in
eukaryotic microorganisms are induced by stressful circumstances such as
overcrowding, resource depletion, and DNA damaging conditions. This
suggests that these sexual processes are adaptations for dealing with stress,
particularly stress that causes DNA damage. In bacteria, these stresses induce
an altered physiologic state, termed competence, that allows active take-up of
DNA from a donor bacterium and the integration of this DNA into the
recipient genome (see Natural competence) allowing recombinational repair
of the recipients’ damaged DNA.
• If environmental stresses leading to DNA damage were a persistent challenge
to the survival of early microorganisms, then selection would likely have been
continuous through the prokaryote to eukaryote transition, and adaptative
adjustments would have followed a course in which bacterial transformation
or archaeal DNA transfer naturally gave rise to sexual reproduction in
eukaryotes.
 Evidences or Theories:
Virus-like RNA-based origin
• Sex might also have been present even earlier, in the hypothesized
RNA world that preceded DNA cellular life forms. One proposed
origin of sex in the RNA world was based on the type of sexual
interaction that is known to occur in extant single-stranded segmented
RNA viruses, such as influenza virus, and in extant double-stranded
segmented RNA viruses such as reovirus.
• Exposure to conditions that cause RNA damage could have led to
blockage of replication and death of these early RNA life forms. Sex
would have allowed re-assortment of segments between two
individuals with damaged RNA, permitting undamaged combinations
of RNA segments to come together, thus allowing survival. Such a
regeneration phenomenon, known as multiplicity reactivation, occurs
in influenza virus and reovirus.
 Evidences or Theories:
Parasitic DNA elements
• Another theory is that sexual reproduction originated from selfish
parasitic genetic elements that exchange genetic material (that is: copies
of their own genome) for their transmission and propagation. In some
organisms, sexual reproduction has been shown to enhance the spread of
parasitic genetic elements (e.g. yeast, filamentous fungi).
• Bacterial conjugation is a form of genetic exchange that some sources
describe as "sex", but technically is not a form of reproduction, even
though it is a form of horizontal gene transfer. However, it does support
the "selfish gene" part theory, since the gene itself is propagated through
the F-plasmid.
• A similar origin of sexual reproduction is proposed to have evolved in
ancient haloarchaea as a combination of two independent processes:
jumping genes and plasmid swapping.
 Evidences or Theories:
Partial Predation
• Another theory is that sex evolved as a form of
cannibalism: One primitive organism ate
another one, but instead of completely
digesting it, some of the eaten organism's DNA
was incorporated into the DNA of the eater.
 Evidences or Theories:
Vaccination-Like Process
• Sex may also be derived from another prokaryotic process. A
comprehensive theory called "origin of sex as vaccination" proposes
that eukaryan sex-as-syngamy (fusion sex) arose from prokaryan
unilateral sex-as-infection, when infected hosts began swapping
nuclearised genomes containing coevolved, vertically transmitted
symbionts that provided protection against horizontal superinfection
by other, more virulent symbionts.
• Consequently, sex-as-meiosis (fission sex) would evolve as a host
strategy for uncoupling from (and thereby render impotent) the
acquired symbiotic/parasitic genes.
Developmental Patterns
 Developmental Patterns
The process by which during development cells become organized in the
embryo is called pattern formation.
All embryos of a given species have a similar structure or body plan. How
does occur? during development each cell must differentiate according to its
position in the embryo, so that the "correct" cell types arise in the correct
place. In other words, cells must know where they are in relation to other
cells in the embryo. This is achieved by giving each cells a positional value
in relation to the principle embryonic axes. Regional
specification describes any mechanism that tells a cell where it is in relation
to other cells in the embryo, so that it can behave in a manner appropriate
for its position. Regional specification is essential for pattern formation.
 Developmental Patterns among
the Metazoa
• We can see that metazoans belong to one of three major
branches:
– Diploblasts
– Protostomes, and
– Deuterostomes.
 Development in Sponges
Sponges develop in a manner so different from that of any other animal
group that some taxonomists do not consider them metazoans at all, and call
them “parazoans.” A sponge has three major types of somatic cells, but one
of these, the archeocyte, can differentiate into all the other cell types in the
body. Individual cells of a sponge passed through a sieve can reaggregate to
form new sponges. Moreover, in some instances, such reaggregation is
species-specific: if individual sponge cells from two different species are
mixed together, each of the sponges that re-forms contains cells from only
one species. In these cases, it is thought that the motile archeocytes collect
cells from their own species and not from others. Sponges contain no
mesoderm, so the Porifera have no true organ systems; nor do they have a
digestive tube or circulatory system, nerves, or muscles. Thus, even though
they pass through an embryonic and a larval stage, sponges are very unlike
most metazoans. However, sponges do share many features of development
(including gene regulatory proteins and signaling cascades) with all the other
animal phyla, suggesting that they share a common origin.
 Development in Diploblasts
• Diploblasts
• Diploblastic animals are those who have ectoderm and
endoderm, but no true mesoderm. These include the cnidarians
(jellyfish and hydras) and the ctenophores (comb jellies).
• Cnidrians and ctenophores constitute the Radiata, so called
because they have radial symmetry, like that of a tube or a
wheel. In these animals, the mesoderm is rudimentary,
consisting of sparsely scattered cells in a gelatinous matrix.
 Protostomes and deuterostomes
• Most metazoans have bilateral symmetry and three germ
layers. The animals of these phyla, known collectively as the
Bilatera, are classified as either protostomes or deuterostomes.
All Bilateria are thought to have descended from a primitive
type of flatworm. These flatworms were the first to have a true
mesoderm (although it was not hollowed out to form a body
cavity), and they may have resembled the larvae of certain
contemporary coelenterates.
 Development in Protostomes
• There are two divisions of bilaterian phyla, the protostomes and the
deuterostomes. Protostomes (Greek, “mouth first”), which include the
mollusc, arthropod, and worm phyla, are so called because the mouth is
formed first, at or near the opening to the gut, which is produced during
gastrulation. The anus forms later at another location. The coelom, or
body cavity, of these animals forms from the hollowing out of a
previously solid cord of mesodermal cells.
• There are two major branches of the protostomes.
The ecdysozoa includes those animals that molt. Its major constituent is
Arthropoda, a phylum containing insects, arachnids, mites, crustaceans,
and millipedes. The second major group of protostomes are
the lophotrochozoa. They are characterized by a common type of
cleavage (spiral), a common larval form, and a distinctive feeding
apparatus. These phyla include annelids, molluscs, and flatworms.
 Development in deuterostomes
• Phyla in the deuterostome lineage include the chordates and
echinoderms. Although it may seem strange to classify humans, fish,
and frogs in the same group as starfish and sea urchins, certain
embryological features stress this kinship. First, in deuterostomes
(“mouth second”), the mouth opening is formed after the anal
opening. Also, whereas protostomes generally form their body
cavities by hollowing out a solid mesodermal block
(schizocoelous formation of the body cavity), most deuterostomes
form their body cavities from mesodermal pouches extending from
the gut (enterocoelous formation of the body cavity). It should be
mentioned that there are many exceptions to these generalizations
 Development in Amniote Egg
The evolution of organisms depends on inherited changes in their
development. One of the greatest evolutionary advances—the amniote egg—
occurred among the deuterostomes. This type of egg, exemplified by that of a
chicken, is thought to have originated in the amphibian ancestors of reptiles
about 255 million years ago. The amniote egg allowed vertebrates to roam on
land, far from existing ponds. Whereas most amphibians must return to water
to lay their eggs, the amniote egg carries its own water and food supplies. It is
fertilized internally and contains yolk to nourish the developing embryo.
Moreover, the amniote egg contains four sacs: the yolk sac, which stores
nutritive proteins; the amnion, which contains the fluid bathing the embryo;
the allantois, in which waste materials from embryonic metabolism collect;
and the chorion, which interacts with the outside environment, selectively
allowing materials to reach the embryo.† The entire structure is encased in a
shell that allows the diffusion of oxygen but is hard enough to protect the
embryo from environmental assaults and dehydration. A similar development
of egg casings enabled arthropods to be the first terrestrial invertebrates.
Thus, the final crossing of the boundary between water and land occurred
with the modification of the earliest stage in development: the egg
 ..

Diagram of the amniote egg of the chick, showing the membranes enfolding the 7-day chick
embryo. The yolk is eventually surrounded by the yolk sac, which allows the entry of
nutrients into the blood vessels. The chorion is derived in part from the ectoderm and
extends from the embryo to the shell (where it will exchange oxygen and carbon dioxide and
absorb calcium from the shell). The amnion provides the fluid medium in which the embryo
grows, and the allantois collects nitrogenous wastes that would be dangerous to the embryo.
Eventually the endoderm becomes the gut tube and encircles the yolk.