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Faculty of Science, School of Sciences, Natabua Campus Lautoka BIO706 Embryology Lecture 1: History of Embryology Developmental Biology The past is full of ideas about how organisms developed and where they came from? These range from pure magic To cultural myths To flights of fantasy And, in recent times, To observational and experimentally based inquiry Questions? Where are the plans or instructions for development? How are they interpreted and used? Where are the raw materials for development? How did those raw materials get to the embryo? How are they used? How do different cells and tissues know what to become? How do cells migrate? How do they know where to go? How is neural circuitry established? How do axons find their targets? Etc., etc, etc. One important thing to keep in mind, Development DOES NOT happen by magic. Information and mechanisms at the cellular and/or molecular levels are needed to accomplish development Developmental Biology is Defined by • • • • • • Differentiation Morphogenesis Growth Reproduction Evolution Environmental Integration Developmental Biology is Defined by • • • • • • Differentiation Morphogenesis Growth Reproduction Evolution Environmental Integration What goes on as an embryo develops from a single, pluripotent cell into a complete organism? Majority of developmental systems are multicellular Single-celled systems used to study cellular differentiation Developmental Biology is Defined by • • • • • • Differentiation Morphogenesis Growth Reproduction Evolution Environmental Integration Cell movements & tissue reorganizations How do differentiated cells move and reorganize during development to generate functional organisms? Major point of study for classical embryology Developmental Biology is Defined by • • • • • • Differentiation Morphogenesis Growth Reproduction Evolution Environmental Integration What are the limiting factors for cell division Allometric growth - varying rates of mitosis in regions of the embryo Isometric growth- equal rates of mitosis throughout embryo Developmental Biology is Defined by • Differentiation • Morphogenesis • Growth • Reproduction • Evolution • Environmental Integration The germplasm Gametogenesis formation and maintenance of pluripotent, haploid germ cells (gametes) Recognition & interaction of gametes at fertilization Developmental Biology is Defined by Survivability of evolutionary • Differentiation mutations is limited by • Morphogenesis restraints of embryogenesis • Growth Evolutionary changes must • Reproduction be small changes for the • Evolution embryo • Environmental What are the evolutionary relationships between Integration groups of organisms Developmental Biology is Defined by • • • • • • Differentiation Morphogenesis Growth Reproduction Evolution Environmental Integration Early development of many organisms is influenced by environmental cues Examples: color/shape of butterfly larvae (caterpillars) Early Embryologist Many of the ancient Greek Philosophers were interested in embryology Democritus (ca. 455-370BC): the sex of an individual is determined by the origin of sperm; Male arise from the right testicle and female from left. This hypothesis modified by Phythagoras, Hippocartes and Galen, however gender bias was always evident, For science was the privilege of men; Philosophers mostly positioned female between Men and Animals, So males were supposed to originated from stronger sperm of right testicle. Early Embryologist Greek Philosopher Aristotle in Generation of Animals ca. 350 BC Suggested the different ways that animals are born From Egg: (Oviparity Eg. Birds, Frogs and Most Invertebrates) From Life Birth: (Viviparity Eg. In placental animals and some fish) By production of an egg that hatches inside the body: (Ovoviviparity Eg. Certain reptiles and sharkes) Aristotle considered two basic developmental questions: Do all parts of a developing organism come into existence together and simply grow larger? or Is development a stepwise process characterized by progressive organization and an increase in complexity? Preformation versus Epigenesis Preformation – The organism is preformed as a complete miniature structure in the sperm or the egg and simply grows larger as it develops. This means that the first reproducing human would have had to have all succeeding generations within itself.. Epigenesis The organism develops in a stepwise fashion from an unorganized state. Aristotle believed that the embryo was formed from the menstrual blood as a result of that blood’s interaction with a male factor, called the male dynamic, that was present in the semen. His observations supported the concept of epigenesis, though this term would not be used until the mid to late 17th century. Greek Philosopher Aristotle noted two major pattern of cell division in early development Holoblastic cleavage (Entire egg is divided into progressively smaller cells E.g. Frogs, Mammals) Meroblastic pattern ( In which only that part of egg destined to become the embryo proper divides, with the remainder serving nutritive purpose E.g. Birds) Mid 17th century - 18th century Epigenesis vs Preformation - a matter for debate Preformationists Spermists vs ovists Jan Swammerdam ~ 1672 Spermists - contended Nicholas Malebranche ~1673 that sperm contained the new individual in Nicolas Hartsoeker ~ 1694 miniature and only Charles Bonnet ~ 1762 nourished in the ovum Ovists- thought the same Epigeneticists way about the ovum and William Harvey ~ 1651 stimulated only in the Rene Descartes ~ 1664 seminal fluid. Marcello Malpighi ~ 1672 Pierre Maupertuis ~ 1745 Preformationists . SPONTANEOUS GENERATION Recipe for bees: Kill a young bull, and bury it in an upright position so that its horns protrude from the ground. After a month, a swarm of bees will fly out of the corpse. Preformationists Jan Baptista van Helmont’s recipe for mice: Place a dirty shirt or some rags in an open pot or barrel containing a few grains of wheat or some wheat bran, and in 21 days, mice will appear. There will be adult males and females present, and they will be capable of mating and reproducing more mice. Preformationists Jan Swammerdam, ~1672 : 17th century Dutch microscopist Debunked “Spontaneous generation” using meticulous dissections and careful experimentation. Thought to have originated the idea of Preformation Greatest contribution to science demonstrated that in insect development, the same organism persists through various stages, i.e. larva, pupa, juvenile, adult. Preformationists Marcelo Malpighi, ~1672 Professor of medicine and personal physician to Pope Innocent XII Early microscopist One of the first scientists to study structures such as the lungs, kidneys, spleen, brain, and skin Because of the importance of his early work a number of anatomical structures still bear his name - Malpighian corpucles (renal corpuscle) in kidney, Malpighian layer in epidermis of skin Malpighian tubules in insects Marcello Malpighi - did not believe what was right before his eyes when examining chicken development: identify the neural groove, the somites and circulation of blood in arteries and veins to and from yolk with the microscope. Preformationists Nicolas Hartsoeker, ~1694 Dutch mathematician and physicist Invented the screw-barrel microscope Co-discoverer of sperm and claimed to be able to see a tiny human –a homunculus- in the head of each human sperm.. The homunculus Preformationists Charles Bonnet, ~1762 Swiss lawyer, naturalist, philosopher Ovist - from studies of parthenogenesis in Daphnia - felt that the theory of preformation was “…one of the greatest triumphs of rational thought over sensual conviction.” In Philosophical Palingests, or Ideas on the Past and Future - argued that females carry within them all future generations in miniature form. Comment on the preformation paradox: “…it is always possible, by adding zeros, to crush the imagination under the weight of numbers.” Epigeneticists William Harvey, ~1651 Physician to King Charles I of England In 1628 Harvey published An Anatomical Study of the Motion of the Heart and of the Blood in Animals which explained how blood was pumped from the heart throughout the body, then returned to the heart - recirculation of blood. Also published Essays on the Generation of Animals - considered the foundation for modern embryology Epigeneticists Rene Descartes, ~1664 Philosopher, physicist, physiologist and mathematician – famous in all. Considered one of the most important and influential thinkers in human history. Cogito ergo sum - “I think, therefore I am.” “If you would be a real seeker after truth, it is necessary that at least once in your life you doubt, as far as possible, all things.” Epigeneticists Pierre Maupertuis, ~1745 French mathematician and biologist In Essai de cosmologie - introduced the theory of survival of the fittest. Argued that preformation could not account for hybrids or “congenital monsters” Proposed that the embryo goes through a number of distinct developmental stages. What’s the correct answer, epigenesis or preformation? Neither is totally correct as originally stated; however, there’s some truth to both, though not in the context of 17th and 18th century thought. Epigenesis - in that development does occur gradually in a stepwise progression moving from a single cell to a multicellular organism of increasing complexity (however, development does not start out from an unorganized state, the zygote is highly organized at the cellular and molecular level). Preformation - in that the instructions for development are present “preformed” in the zygote - genes (DNA). The preformation/epigenesis issue was the subject of vigorous debate throughout the 18th century. But the problem could not be resolved until one of the great advances in biology had taken place-the recognition that living things, including embryos, were composed of cells. Cell theory of development The cell theory developed between 1820 and 1880 by, the German botanist Matthias Schleiden and the physiologist Theodor Schwann. It was at last recognized that all living organisms consist of cells, which are the basic units of life, and which arise only by division from other cells. Multicellular organisms such as animals and plants could then be viewed as communities of cells. Development could not therefore be based on preformation but must be epigenetic, An important advance was the proposal by the 19th century German biologist August Weismann that the offspring does not inherit its characteristics from the body (the soma) of the parent but only from the germ cells-egg and sperm-and that the germ cells are not influenced by the body that bears them. Weismann thus drew a fundamental distinction between germ cells and somatic cells or body cells The distinction between germ cells and somatic cells. In each generation germ cells give rise to both somatic cells and germ cells, but inheritance is through the germ cells only In 19th century an experiment on sea urchin eggs showed that after fertilization the egg contains two nuclei, which eventually fuse; one of these nuclei belongs to the egg while the other comes from the sperm. Fertilization therefore results in an egg carrying a nucleus with contributions from both parents, and it was concluded that the cell nucleus must contain the physical basis of heredity. The climax of this line of research was the eventual demonstration, toward the end of the 19th century, that the chromosomes within the nucleus of the zygote (the fertilized egg) are derived in equal numbers from the two parental nuclei, and the recognition that this provided a physical basis for the transmission of genetic characters according to laws developed by the Austrian botanist and monk Gregor John Mendel. The coming together of genetics and development Genetics was seen as the study of the transmission of hereditary elements from generation to generation Embryology was the study of how an individual organism develops and, in particular, how cells in the early embryo became different from each other. Danish botanist Wilhelm Johannsen in 1909 put forward a concept to link genetics and embryology was distinction between genotype and phenotype The genetic endowment of an organismthe genetic information it acquires from its parents-is the genotype. Its visible appearance, internal structure, and biochemistry at any stage of development is the phenotype. The discovery in the 1940s that genes encode proteins was a major turning point. As it was already clear that the properties of a cell are determined by the proteins it contains, the fundamental role of genes in development could at last be appreciated. By controlling which proteins were made in a cell, genes could control the changes in cell properties and behavior that occurred during development. Questions are welcome