Adopted from: Dr Neil Vargesson DevBiol The objective of this course 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 behaviour and the formation of an embryo. DevBiol
Biologists study development for different reasons.
One reason is to understand how a single cell (the zygote) can produce the variety of body parts in an organism.
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 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) 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! Preformation is the concept of a miniature adult being present in the sperm or egg, waiting to unfold.
Some claimed they could see a miniature adult in the egg or sperm (homunculus theory). A young animal is merely unfolding the structures that are already there. Early Concepts: Preformation Versus Epigenesis Epigenesis is where new structures arose progressively through a number of stages Aristotle 384BC - 322BC
In 1759, Kaspar Friederich Wolff showed there was no preformed chick in the early egg.
-Undifferentiated granular material became arranged into layers.
-The layers thickened, thinned, and folded to produce the embryo.
-Epigenesis is the concept that the embryo contains building materials that are assembled progressively. The Cell Theory - Schleiden and Schwann (1838) Organisms are composed of cells, the basic units of life. Both animals and plants are multicellular composites that arise from a single cell, thus, development is epigenetic. The fertilised egg is a specialised SINGLE cell (zygote). Only the germ cells (egg and sperm) pass on characteristics on to the offspring. Sea urchin expts showed two nuclei in 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 Induction A cell, or tissue, directs the development of another.
The importance of induction and other cell-cell interactions was demonstrated dramatically by Hans Spemann and Hilde Mangold in 1924. 1940s and 1960s o Genes shown to encode proteins and later such proteins influence other genes. o This finding finally linked genes with embryology! o Since then, a huge explosion in genes/embryology. We will discuss some examples as this course progresses.
Today A major goal of developmental biology is to understand how genes control embryonic development. Model Organisms
Genetic control of development is studied in a variety of animal models. These are chosen for historical reasons, as well as ease of study and biological interest.
Each has advantages and disadvantages and most labs work with more than one model organism, in order to gain the best insights. Impact on Society IVF Teratology Dietary advice Chromosomal basis of birth defects Screen for genetic mutations causing birth defects Future Impact Understanding developmental mechanisms will aid in designing therapies for cancer, diabetes etc Animal models of human disease Regeneration Prenatal screening Organ harvesting/transplantation Embryonic stem cells Cloning Next Common Features in Development Common Features of Development I
Adopted from: Dr Neil Vargesson Typical animal life cycle and stages Animal pole Vegetal pole anterior ventral dorsal Medio lateral ventral dorsal Animal pole Vegetal pole posterior 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 and the next lecture describe these common features or principles
All (or nearly all) animal embryos have the following features:
1. Cleavage/cell division: the process by which a single-celled 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 eg: A-P axis
4. Cell differentiation - where different sorts of cells arise - more than 200 types in a vertebrate body.
5. Growth - increase in size.
Eg: 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 behaviour. Development is a Series of Progressive Changes
This begins when a fertilised egg divides mitotically.
Specialisation 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 chromosone number Fertilisation - diploid chromosone number
Maternal and paternal chromosones are the blueprint for a new individual Meiosis leads to the haploid number of chromosones Meiosis ensures no two offspring of parents are genetically identical Spermatogenesis Oogenesis Fertilisation
Fertilisation is the union of male and female gametes.
Fertilisation provides for recombination of paternal and maternal genes, restoring the diploid number.
Fertilisation activates the egg to begin development.
Fertilisation requires the Acrosome reaction Prevention of Polyspermy
Polyspermy, the entry of more than one sperm, would cause a triploid nucleus.
Important changes in the egg surface block entrance to any additional sperm.
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 Different animal groups undergo different amounts of cleavage.
Eg: 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.
The types of cleavage a - when cleavage complete and egg is divided into blastomeres. b - eg chick where cytoplasm located at animal pole and only this region cleaves c - eg: insects and crustaceans d - eg: mammals e - eg: molluscs, worms Human cleavage and blastula formation 30hr 3 days 4.5-5 days 6 days 48hr 4 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 The bilaminar disc (approx 9 days) Blastocyst Embryo Foetus 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 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 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 Next lecture: Common features of development II - review of this lecture, then onto Axes, symmetry, morphogenetic processes, growth, death and the role of the genes.