Introduction to Animal Embryonic Development

The objective of this lecture is to introduce you to the processes underlying development. The big picture is to understand how animals develop from a single cell to a multicellular organism. We will explore the mechanisms controlling cell behavior and the formation of an embryo. There will be much emphasis on model genetic organisms.

What Can We Learn from Development?

Biologists reasons. study development for different

1) One reason is to understand how a single cell (the zygote) can produce the variety of body parts in an organism. 2) Another reason is the search for commonalities among organisms. The division of the 32 multicellular animal phyla into groups is simplified by an understanding of their development.

Why?

Why study development?

- Development of structures. - How normality arises. - How abnormality ensues. - Positioning of adult structures. - Understand that several tissues form at same time, requiring same genes (e. g. HOS- Holt-Oram Syndrome) thus a defect in one tissue can indicate defects elsewhere

Holt-Oram Syndrome - heart/hand defects

Atrial septation defects Range of hand abnormalities

Phenotype due to mutation in one gene required at time both structures develop thus knowing relationship of structure/organ development will aid in diagnoses!

The Cell Theory - Schleiden and Schwann (1838)

1. Organisms are composed of cells, the basic units of life. 2. Both animals and plants are multicellular composites that arise from a single cell, thus, development is epigenetic. 3. The fertilized egg is a specialized SINGLE cell (zygote). 4. Only the germ cells (egg and sperm) pass on characteristics on to the offspring. 5. Sea urchin experiments showed two nuclei in one egg - 1 from egg and 1 from sperm.

Determination and Regulation

Once recognized that cells in the embryo arose by cell division - the question then emerged how do cells become different from one another?

Determination: cell nuclei of the early embryo contains determinants, that are distributed unequally to daughter cells & control cells future (mosaic).

Regulation: embryo can form normally even if parts of the embryo are missing, implying cells interact with each other.

Example of regulation in embryonic development

Development from a single cell into a multicellular organism is the most complicated fate a single cell can undergo.

Development is essentially the emergence of organized structures from an initially simple group of cells.

Yet, only a few basic principles are needed (for the majority of animal organisms to form).

This lecture describe these common features or principles

Most animal embryos have the following features:

1. Cleavage/cell division: the process by which a singlecelled zygote divides into smaller units, blastomeres. 2. Morphogenesis - at various developmental time-points embryos undergo changes on 3D form - the most striking changes in form are (i) gastrulation: the process by which the embryo forms different tissue layers from which future organs will be built; (ii) neurulation; (iii) coelom formation. 3. Regional specification or pattern formation - where pattern appears in a previously similar population of cells and initially involves laying down of a body plan e. g.: A-P axis.

4. Cell differentiation - where different sorts of cells arise - more than 200 types of cells in a vertebrate body.

5. Growth - increase in size.

E. g.: pattern formation in early development specifies differences between cells that lead to changes in form, cell differentiation, and growth.

It is gene expression that controls all these processes, changing patterns of gene expression during early development change cell identities, giving rise to their future behavior.

Development is a series of progressive changes

This begins when a fertilized egg divides mitotically. Specialization occurs as a hierarchy of developmental decisions. Cell types do not unfold but arise from conditions created in preceding stages. Interactions become increasingly restrictive; each stage limits developmental fate. With each new stage, cells lose the option to become something differentit becomes determined. Both cytoplasmic localization and induction cause this feature.

Gametes
Male germ cell sperm or spermatozoa Female germ cell secondary oocyte Meiosis - haploid chromosome number. Fertilization - diploid chromosome number. Maternal and paternal chromosomes are the blueprint for a new individual.

Meiosis leads to the haploid number of chromosomes

Meiosis ensures no two offspring of parents are genetically identical

Spermatogenesis

Oogenesis

Fertilization
Fertilization is the union of male and female gametes. Fertilization provides for recombination of paternal and maternal genes, restoring the diploid number. Fertilization activates development. the egg to begin

Fertilization requires the acrosome reaction

Prevention of Polyspermy
Important changes in the egg surface block entrance to any additional sperm. Polyspermy, the entry of more than one sperm, would cause a triploid nucleus. In the sea urchin, an electrical potential rapidly spreads across the membrane; this is the fast block. This is followed by the cortical reaction, where enzymatic and metabolic changes trigger cytoplasmic rearrangements.

Fusion of Pronuclei and Egg Activation

After sperm and egg membranes have fused, the sperm disconnects from its flagellum. The enlarged sperm nucleus is the male pronucleus and migrates inward to contact the female pronucleus. Fusion forms a diploid nucleus. Nuclear fusion takes 12 minutes in sea urchins; about 12 hours in mammals. The fertilised egg is now properly called a zygote.

Fertilisation initiates reorganisation of cytoplasm and repositions determinants that begin development and cleavage.

A typical zygote is small, spherical and polarized along vertical axis, establishes the direction of cleavage and differentiation. Upper hemisphere = animal hemisphere (or pole) Lower hemisphere = vegetal hemisphere (or pole) and is rich in yolk. Early cell divisions are called cleavages The embryo undergoes cleavage to convert the large cytoplasmic mass into small maneuverable cells (blastomeres). No cell growth occurs, only subdivision until cells reach regular somatic cell size.

Xenopus cleavage (sub-saharan frogs)

Different animal groups amounts of cleavage.

undergo

different

E. g.: At the end of cleavage, polychaete worms have 1000 cells, amphioxus has 9000, and frogs have 700,000.

Different animal groups use different types of cleavage to obtain the ball of cells (BLASTULA) that will eventually produce an adult organism.

a when cleavage complete and egg is divided into blastomeres. b e. g. : chick where cytoplasm located at animal pole and only this region cleaves c e. g. : insects and crustaceans d e. g. : mammals e e. g. : molluscs, worms

30 hr

48 hr

3 days

4 days

4.5-5 days

6 days

Gastrulation
After blastula formation, almost all animal embryos undergo GASTRULATION. This process varies in different species, but is essentially produces the same outcome.

A phase of cell movements occurs that converts the ball (mammal) or sheet of cells (bird/fish) into a three layered structure = GASTRULA.

These three layers are the GERM LAYERS. Ectoderm, mesoderm and endoderm.

The bilaminar disc (approx 9 days)

Formation of a trilaminar embryo: Gastrulation

Day 15/16

Gastrulation and germ layer formation

Neurulation
Formation of a neural tube develops into brain and spinal cord.

Slight differences between species, but essentially produces the CNS.

Organogenesis Begins With Development of the Nervous System

The nervous system is the first organ system to develop. a) The notochord grows and induces overlying ectoderm to form the neural plate. b) Cells of the neural plate fold to form the neural groove and the surrounding neural folds.

c) The neural folds fuse, forming a hollow neural tube. The anterior portion forms the brain; the rest forms the spinal cord. Right, neural fold formation in the human embryo

After completion of the major morphogenetic processes, most types of animal embryo have a body plan, but the body is yet to differentiate. This is also known as the phylotypic stage

Blastocyst

Embryo

Foetus

Early in development embryos of different species look very similar.

Fish, chick, mouse used as model genetic systems to study human diseases as genes used for development are the same

 Overview: A Body-Building Plan for Animals. It is difficult to imagine: That each of us began life as a single cell, a zygote

 The question of how a zygote becomes an animal?

Has been asked for centuries.

As recently as the 18th century.

The prevailing theory was a notion called preformation.

Preformation is the idea that the egg or sperm contains an embryo

A preformed miniature infant, or homunculus, that simply becomes larger during development

An organisms development: Is determined by the genome of the zygote and by differences that arise between early embryonic cells Cell differentiation: Is the specialization of cells in their structure and function Morphogenesis: Is the process by which an animal takes shape

 After fertilization, embryonic development proceeds through cleavage, gastrulation, and organogenesis. Important events regulating development: Occur during fertilization and each of the three successive stages that build the animals body.

 The main function of fertilization: Is to bring the haploid nuclei of sperm and egg together to form a diploid zygote. Contact of the sperm with the eggs surface: Initiates metabolic reactions within the egg that trigger the onset of embryonic development.

Spiral Cleavage in an Annelid Embryo

 Radial cleavage Very similar to the sea star Deuterostomes

blastula

blastula

gastrulas

 A human embryo at approximately 68 weeks after conception

Shows the development of distinctive features

1 mm

Organ Development
Begins during the first trimester: Gastrulation occurs during the 2nd and 3rd weeks, followed by neurulation (formation of the neural tube) The heart beats spontaneously after 3.5 weeks After the first two months of development, the products of conception are called a fetus

Week 5

 At the end of the first trimester (first 3 months of development): Fetus can be recognized as a human ~56 mm long, and ~14 g The sexes can be differentiated Ears, eyes becoming welldeveloped, Skeleton starting to develop Notochord replaced with the developing vertebral column Moves, breathes, makes sucking motions with thumb

33-day embryo measuring 7 x 3.2mm.

Human embryo at 40 days.

Human Fetus at Ten Weeks

7-week fetus in uterus

10-week fetus in uterus

 In the second trimester:

Fetus moves freely Heart can be heard with a stethoscope Quickening movements felt by the mother If born at 24 weeks, the fetus has a 50% chance of survival Brain not yet able to support breathing Kidneys and lungs are immature

20-week fetus in uterus

 In the third trimester: Fetus grows rapidly Final differentiation of organs and tissues Grasping and sucking reflexes active If born before 37 wks, is considered premature, but has a good chance of surviving if born after 30 weeks Full-term baby weighs about 5000 g and is 52 cm long

Chicken embryo, 43 hours

Chicken embryo, 60 hours

Chicken embryo, 8 days

Chick embryo, 9 days