SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. Origin of life theories & Evolutionary theory “Oh, in truth, at that moment I deprived the culture medium of the only thing not given to man to produce (… life).” L. Pasteur, 1864 There is more than 2 million different forms of life on planet earth today and we have to assume (based on fossil records and other found traces of life) that there existed an even greater number of different life forms in the millions of years before us Even though modern science is still not able to explain how life got on this planet, it offers a series of intellectually very interesting theories, or origin of life theories, which are strongly founded on important experimental observations and which will be laid out in some detail in this chapter A significant body of scientific evidence shows that life is on earth since approximately 1 billion years and strongly supports the idea that primitive life forms may have existed on earth as early as 3.5 - 4 billion years ago during this tremendously vast period of time life may have evolved from primitive precellular structures and out of a “molecular broth” (see “theories on the origin of life” in the sections below) into millions of genetically and morphologically different species which call this fascinating blue planet home today 2 million different species are known to live on Earth and they populate almost all regions on this life-supporting planet early biologists, most prominently Karl von Linne, started to classify the huge variety of species into so-called taxa and introduced the first taxonomic system into biology; this early taxonomic system was primarily based on morphological characteristics, i.e. existence or absence of certain body features; body shape, etc. in biological organisms with the discovery of DNA, the isolation of proteins and other biomolecules, together with the introduction of a plethora of molecular biological techniques e.g. DNA sequencing, protein sequencing, chromosome analysis, NMR analysis, karyotyping, etc. and the establishment of comprehensive DNA and protein data bases, modern biologists started to classify organisms based on their molecular similarities, e.g. DNA homologies, protein sequence similarities, rather than due to their morphological characteristics - most of the existing “traditional” phylogenetic trees and relationships are challenged these days by new findings or these new approaches today, all existing species on our planet are classified into the so-called five major kingdoms of organisms for each of these 5 kingdoms a so-called phylogenetic tree has been developed on the basis of morphological and genetic features of each of its members 1 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. the five kingdoms of living organisms on planet Earth are: 1. 2. 3. 4. 5. Bacteria Protista Fungi Planta Animalia As different as all these different forms of life might be on the tissue land organ level and regarding their intricate body morphologies, “deep down” they all share a plethora of structures and molecules on the cellular and molecular level, such as ATP, DNA, RNA, proteins made up from amino acids, and phospholipids-made cell membranes and organelles This modern day information is important in the critical and fair evaluation of contemporary and also future theories about one of the most challenging questions to be answered in the biological science; the question about where life came from, how it established itself on planet earth and how it turned itself into one of the great wonderful success stories in this known universe with more than 2 million different life forms today Origin of Life Theories Indeed, the question of how and when life arose on planet Earth has fueled the scientific discussion and our imagination ever since the dawn of human mankind, the evolution of the human brain and conscience the origin of life on Earth has been dated back to 3.5 – 4.0 billion years; the earliest, morphologically primitive life forms appear in certain rock formations during that time period - various fossil records of the Precambrian period (4,600 – 600 Mio. years ago) have been found in certain geological rock formations (see Images below) - the images below show fossilized cyanobacterial colonies, so-called stromatolithes (left panel) as well as fossilized images of filamentous (top right) and colonial (top bottom) photosynthetic cyanobacteria (e.g. Gloeocapsomorpha), which have been found in 1- 3 billion year old Archean rock formations, e.g. in West Australia (Bitter springs chert) they were identifiable by chemical breakdown products of their photosynthesis pigments (“chemical fossils”) - further evidence comes in form of Pisolites, which are small concentrically layered structures of fossilized bacteria 2 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. filamentous cyanobacterium Palaeolyngbya Layered stromatolite (= calcium carbonate precipitate) colonial chroococcalean cyanobacterium ( 850 mya) but how and under which circumstances did already complex organized and structured life get there? Speculation about the true origin of life and where life comes from has a long human history and can be traced back to the early human civilizations (for an overview see the Graphic below) 3 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. Origin of Life Theories & Milestone Discoveries Molecular Biology Organic Chemistry Alchemy Microbiology Graphic©E.Schmid/2003 Ancient Times 1700 1800 1861 1828 “life arose from non-living matter” (Aristotle) “Molecular Evolution Theory” (M. Eigen, L. Orgel) Proteinoid Microspheres (S. Fox) “Phlogiston Theory” (J. Becher, G. Stahl) “Vitalism Theory” (G. Stahl) 1850 Synthesis of glycine (W. Loeb, Klages, et al.) Chemical synthesis of urea (F. Woehler) from silver cyanite & Ammonium chloride vis vitalis exists 1900 Synthesis of alanine (Strecker) Synthesis of sugar (Butlerov) 1970s 2000 1938 1953 1980s Miller-Urey Experiment (S. Miller) “RNA world theory” Ribozymes (T. Czech) “life begets life” (Virchov, Pasteur) “Anoxic Early Earth Atmosphere Theory” (A.I. Oparin, J.B.S. Haldane) in aanncciieenntt ttiim meess, the Greek philosopher Aristotle speculated that life arose from nonliving matter - when Aristotle introduced his “spontaneous generation” idea, he together with other thinkers of his time, such as Democritus and Epicurus, had no doubt that life naturally originated from inert, inorganic matter The concept of spontaneous generation was widely accepted and unquestioned far into the middle of the 17th century and was supported by some of the thinkers, teachers and scholastics of the Church, including Saint Basil (329 – 379), Saint Augustine, Saint Thomas of Aquinas as well as naturalists like J.B. van Helmont - since the existence and true nature of seeds, egg cells and molecules has not been discovered yet, it was common believe that flies and maggots form from components of rotten meat by spontaneous generation and that bacteria are directly created form broth components over time In 1668, the Italian physicist Florencia Francisco Redi (1626 – 1697) performed his famous maggot experiment which experimental outcome lead to serious doubts about the correctness of the upheld spontaneous generation idea of his time - he used two glass containers in both of which he brought pieces of meat; while he left the neck of one of the vials open, he blocked the entry into the second vial by placing some loose cotton wool 4 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. - maggots only developed in (and flies only flew out of) the first vial, while no insect development occurred in the second container the maggots must have got there by some inoculation by flies (which as we know today) laid their eggs into the piece of meat Around the 1700s, the Italian physician Antonio Vallisnieri (1661 – 1730) proved that hatched insects from plant galls derive from previously laid eggs into the plant tissue, which later on grew in size and helped to nourish the developing caterpillars of the insect - this careful observation gave a further argument against the popular spontaneous generation theory in the 1177ttthhh cceennttuurryy, the German physicians and university professors J. Becher and G. Stahl develop the so-called “Phlogiston theory” (from the Greek word phlogistos = burning) to explain chemical combustion reactions - Stahl further believed living matter contained a “soul” (vis vitalis) that differed in composition from nonliving matter (“Vitalism theory”) - He and many generations of vitalism supporters after him searched without success to collect the phlogiston and to find proof for the “mysterious life force” in living beings until the m miidd--11880000''ss scientists widely accepted the Vitalism theory and generally believed that organic (= carbon-containing) chemicals could only form through living things and with the help of a mysterious “vital force” in 11882288, the German chemist Friedrich Woehler synthesizes the first organic compound, called urea, from silver cyanite and ammonium chloride, two purely inorganic materials; his ground-breaking chemical experiment disproved the dominating Vitalism theory of his time Lingering doubts on the validity of the experimental design of the experiments which claimed to disprove the spontaneous generation idea were put aside when the great French scientist Louis Pasteur (1822 – 1895) conducted his famous experiment at the Paris Academy of Sciences - after heating the nutritional broth in the bulb of his designed swan neck flask, no bacterial growth and life arose in the open flask after even extended period of time - due to the long curvature of the flask neck, no bacteria or spores were able to reach the broth in the interior of the flask to trigger growth of new life after the lifedestroying heating procedure - the outcome of this elegant experimental setup finally disproved the long upheld spontaneous generation idea “There is no spontaneous generation of life possible under the currently existing life conditions and in our currently existing highly oxidative earth atmosphere of today … “ 5 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. in the llaattee 1199ttthhh cceennttuurryy, LLoouuiiss P Paasstteeuurr and R Ruuddoollff V Viirrcchhoow w concluded from their observations and studies, that life arises only through reproduction of pre-existing life; “life begets life” - there is no spontaneous generation of new biological organisms possible under the current Earth conditions - generation of new life requires unique reproductive structures, e.g. spores, or a special type of cells, i.e. generative cells, such as sperm and egg cells in 11886611, the Russian chemist Butlerov succeeds to chemically synthesize the six carbon-molecule glucose from inorganic precursors Around 11990000, the Swedish scientist Svente Arrhenius (1859 – 1927) proposes and advocates the hypothesis that life did not originate on Earth but was seeded from outer space ( begin of the cosmic ancestry hypothesis) in 11993388, the Russian biochemist A.I. Oparin and the British scientist J.B.S. Haldane suggested that carbon-molecules that served as the basis of life were formed in an early (primordial), oxygen-less (= anoxic) Earth atmosphere which consisted of methane, ammonia, hydrogen, and water vapor “Primordial Anoxic Earth Atmosphere Theory” - according to the core ideas of this theory, the early anoxic earth had a reducing atmosphere which may have favored the spontaneous, reductive build-up (chemical synthesis) of more complex molecules - today, the modern earth atmosphere with its 21% oxygen content is a strongly oxidizing environment which rather favors spontaneous oxidative chemical processes in 11995533, the American biochemist Stanley Miller tested this idea and showed for the first time, that complex, organic compounds, such as amino acids, form spontaneously under controlled lab conditions; the foundation of the “prebiotic broth theory” is laid out his famous “Miller-Urey apparatus” was a closed experimental system which simulated the primordial composition of the oxygen-free atmosphere of the ancient Earth (= primordial earth atmosphere), atmospheric discharges (lightning), volcanic activity and considered the presence of an early ocean made of water (see Figure below) it contained methane, ammonia, water (vapor), and hydrogen which circulated past a source of electric discharges it resulted in the generation of amino acids (alpha-alanine, beta-alanine, aspartic acid, alpha-amino-n-butyric acid, and several other unidentified amino acids) “The prebiotic broth theory is introduced into the origin of life discussions.” 6 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. Simulation of the ancient Earth atmosphere and Earth by Urey & Miller (1950s) in their famous Urey- Miller apparatus lead to the build-up of amino acids and other complex organic molecules in their “pre-biotic soup” Primordial atmosphere simulated Electrical discharges CH4, H2 NH3 simulated Lightning yielded Amino acids, Organic molecules H2O simulated Ancient oceans simulated Volcanic activity/ Lava - different energy sources and proportions of gases used in the same experimental setting even yielded purines, pyrimidines, and carbohydrates: all essential building block molecules for life 7 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. in 11996600, Juan Oro’ experimentally obtains the biomolecule adenine after mixing ammonia (NH3) and hydrocyanic gas with water and heating it to 194oC adenine is the crucial precursor molecule for many important biomolecules, most importantly ATP, DNA, RNA, FAD and NAD in the 11997700ss, Sidney W. Fox demonstrates that amino acids, the building blocks of proteins found in all living things, spontaneously organize themselves into cell-like structures, so-called "protenoid microspheres” or proto-cells (see Figure below); he showed the formation of organic polymers after dripping dilute aqueous solutions of organic monomers on hot sand or clay the proto-cells studied by S. Fox exhibited some of the features of living cells, such as growth, membranous responses, metabolism and transfer of chemical information Spontaneous formation of Proto-cells ( Sidney Fox experiments) Formation of free protocells on the surface of abiotic thermal protein TP9 Protocell Multi-protocells of thermal protein SIU-2 (diameters range from 0.3-0.6 μm) 8 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. in the 11998800ss, the American biochemist Tom Czech showed for the first time that certain RNA molecules have enzymatic activity and work as so-called ribozymes (see Figure below) the idea was developed whereas early life-like processes, e.g. a primitive form of metabolism, my have started in a so-called “RNA-world” with ribozymes as the primordial enzymes on this planet in modern living organisms today, catalytic RNAs or ribozymes still play a significant role in many biological processes ribozymes and catalytic active RNA molecules are known to be required: 1. for the viability of certain plant pathogens 2. for the “RNA business” of the human hepatitis delta virus (HDV) self-cleaving/RNA processing 3. as functional units of ribosomes (23S rRNA) for protein synthesis 4. as functional units of telomerases to protect the chromosomal ends from “replicative erosion” 5. for gene self-splicing processes in the protozoan Tetrahymena Five classes of ribozymes have been described of which the “hammerhead” ribozyme is the best studied one (see detail A of Figure below) Consensus structure (A) and turn-over cycle (B) of a “hammerhead’ ribozyme A B By the 11998800ss, many scientists consider protenoid microspheres with entrapped RNA molecules as proto-cells and therefore the smallest unit of proto-life 9 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. in 11999922, Guenter Waechtershaeuser introduces the “Pressurized iron-sulfur world theory” which suggests a deep ocean hydrothermal origin of a primitive autotrophic metabolism of small carbon molecules, most importantly pyruvate and acetate, in a deep ocean vent environment of earth crust iron sulfides and hot magmatic exhalations (for an overview see Graphic below) on the early, protein-less world, mineral-bound iron and nickel sulfides may have had the function of inorganic catalysts of a primitive metabolic cycle according to this theory, the inorganic iron and nickel sulfides surfaces of deep ocean vent clefts may have served as early catalysts, before they were eventually succeeded by the more effective protein-embedded minerals in form of cofactors of enzymes the mineral-bound iron and nickels sulfates may have mimicked the catalytic activity of acetyl coenzyme A (CoA) synthase, an enormously important iron- and nickel-containing enzyme complex present in modern chemoautotrophic organisms Literature: G. Waechtershaeuser, Science 289: 1307 – 1308 (2000) Chemical reactions & End products in the Iron-Sulfur world cycle Primitive chemical cycle of the deep ocean vent environment? CH3 C CH3 – SH CO2 O Acetic Acid OH Carbon dioxide O HO CH3 Heat CH3 OC C C C H2S HCO2 CH3 CO O O Methyl thioacetate S – CH3 O C – C – CH3 Fe2+, Co, Ni2+ Mineral Heat library CO O Carbon O C – C – CH3 NH3 Ammonia Peptides O Graphic©E.Schmid/2004 adopted with modifications from G. Waechtershaeuser O HO Pyruvic Acid NH2 C – C – CH3 HO Alanine 10 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. In 22000000, G. D. Cody et al. shows experimental evidence that the crucial cell metabolism molecules pyruvic acid, acetic acid and alanine, form from formic acid in the presence of carbon monoxide, nonylmercaptane and iron sulfilde at high pressures (0.2 -200 MPa) and temperatures (100 - 250oC) (see Graphic above) he introduces the idea that important cell metabolism intermediates may have formed through the reaction of alkyl sulfurs and carbon monoxide (CO) with iron sulfide in ancient deep ocean hydrothermal vents (“cold smokers”) he speculates that this chemistry may have provided the critical biomolecules for early, primitive metabolic processes on the prebiotic earth Literature: George D. Cody, et al., Science 289: 1337 – 1340 (2000) “The maxim of the prebiotic broth theory is “order out of chaos”. In contrast, the iron-sulfur world experiments are serial, aimed at long reaction cascades and catalytic feedback (metabolism) from the start. The maxim of the ironsulfur world theory should therefore be “order out of order out of order” G. Waechtershaeuser in 22000033, C. Huber & G. Waechtershaeuser (Technical University Munich, Garching, Germany) showed proof for a primitive peptide cycle with the amino acid phenylalanine under test tube conditions which may have occurred on the primordial world, e.g. in certain deep ocean environments (“black smokers”) the observed peptide cycle had anabolic and catabolic segments resembling a modern metabolic cycle operating in living organisms they showed carbon monoxide (CO)-driven build-up of short peptides from α-amino acids under hot aqueous test tube conditions in the presence of colloidal iron-nickel sulfide [ (Fe,Ni)S ] minerals they also reported the CO- and (Fe,Ni)S ] mineral-dependent degradation (hydrolysis) of a dipeptide (L-Phe- L-Phe) down to amino acid monomers Literature: C. Huber, et al., Science 301: 938 – 940 (2003) in 22000044, the NASA scientist Jason Dworkin (Goddard Space Flight Center, Maryland), introduces the idea that early biological membranes might have been formed inside dense interstellar clouds and may have been seeded on earth via impacting asteroids his idea his based on the experimental results he received with lab-simulated space dust he mixed with water, methanol, ammonia, carbon monoxide in the extreme cold (- 258 oC) under the influence of strong UV irradiation under these experimental set-up conditions he found organic membranes that selfassembled into hollow, spherical structures Even though our modern scientific understanding of one of the greatest mysteries in the universe, the origin of life, is far from being understood, today, the scientific community favors 3 major theories how life may have started out and evolved on our planet 11 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. 1. Prebiotic broth theory & Organic chemical evolution theory The core idea of the prebiotic broth theory is that life slowly originated from inorganic matter in a prebiotic broth under the influence of natural physico-chemical forces either on earth of out there in the universe; according to this theory, carbon-based molecules (once formed through physico-chemical forces) created a molecule-enriched prebiotic broth in which self-assembly eventually formed self-sustaining and stable high molecular weight carbon-structures, such as RNA, proteins and lipids through a rigorous “sorting out mechanism” ( selective pressure & chemical evolution) The core idea of the organic chemical evolution theory assumes that the early life forms evolved by macromolecular build-up from simple carbon precursor molecules by chemical synthesis processes; according to this theory macromolecules assembled spontaneously and evolved in a totally different earth atmosphere than the one we know today and which we breathe; the theory (which is associated with the scientists M. Eigen & L. Orgel) is strongly based on scientific evidence supporting an oxygen-free, ancient, so-called anoxic primordial earth atmosphere (for an overview see table below) and also includes key ideas of the evolutionary theory, such as natural selection and selective pressure Scientific evidence strongly suggests that the early earth atmosphere existed mainly of hydrogen (H2), carbon monoxide (CO), carbon dioxide (CO2), nitrogen gas (N2), methane (CH4), ammonium (NH3) and water vapor Gas composition of the different atmospheres in Earth’ history Chemical Composition of the Earth’ atmosphere today: Nitrogen (N2) 78% Oxygen (O2) 21% Carbon dioxide (CO2) 0.03 % other miscellaneous gases, e.g. H2O the ancient, primordial Earth atmosphere was most likely composed of: Hydrogen (H2) and Helium (He) these two gases are relatively rare on Earth today compared to other places in the universe they were probably lost to space early in Earth's history the second atmosphere establishing on Earth was primarily produced by volcanic outgasing and most likely consisted of: Water (H2O) Carbon dioxide (CO2) Sulfuroxide (SO2) Carbon monoxide (CO) Sulfur (S2) Nitrogen (N2) Hydrogen (H2) Ammonia (NH3) and Methane (CH4) 12 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. this atmosphere most likely contained no free oxygen (O2) at this time, since it is not traceable in gases of modern volcanoes traces of existing oxygen may have been quickly reduced by transition metals as part of the earth crust, such as iron (Fe2+ Fe3+) the chemically reducing (= oxygen-free) earth atmosphere, rather than oxidizing atmosphere favored anabolic (= chemical build-up) reactions and the formation and accumulation of small organic molecules, e.g. amino acids and even more complex polymers in his famous ‘pre-biotic soup’ experiment in the 1950s, S. Miller confirmed the formation of organic molecules under conditions where he simulated the early earth atmosphere conditions in his lab electrical sparks (which simulated atmospheric lightning) triggered the formation of amino acids, nucleotides and other bio-molecules in an atmosphere consisting of water vapor, CH4, NH3 and molecular hydrogen the amount of free O2 in the atmosphere gradually rose from 1 - 10 % during the Proterozoic era due to the activity of photoautotrophic organisms, most importantly the photosynthesizing cyanobacteria and purple bacteria scientists favoring the chemical evolution theory assume that the polymers of life (RNA, peptides) were formed by so-called dehydration synthesis on hot rock or clay formations originating from volcanic activities these regularly flooded clay and rock formations may have served as the first polymerization sites these first genes in form of short strands of RNA laid the basis for reproduction; the most important hallmark of life the first strands of RNA assembled with complementary RNA molecules in the prebionic atmosphere without the assistance of protein structures, but probably with the (catalytic) help of the net charges of transition metals being part of clay or rock surfaces the first rounds of replication occurred these simple replication/assembly processes may have been aided by specially formed and catalytically active RNA molecules, called ribozymes today we now that ribozymes can catalyze RNA splicing and polymerization more recently scientists discovered that the polypeptide-forming cellular complex called ribosome is actually a ribozyme! in the pre-bionic so-called RNA world, RNA build the first genes long before DNA took over this important functional part of life scientists assume that early forms of so-called molecular cooperation must have occurred in chemically unique micro-environments, e.g. electrically charged clay surfaces/cavities on our young planet the first polypeptide chains arose on self-replicating RNA polymers/ribozymes some of these generated polypeptide chains started to act as primitive enzymes and protective/stabilizing proteins the first primitive RNA polymerase may have evolved and started to aid the RNA replication process 13 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. spherical membranes made from hydrated (= flooded) lipid molecules started to enclose the RNA-polypeptide cooperations may have formed the first simple cell-like structures or so-called proto cells after millions of years of repeated rounds of trial and error and millions of probed molecular cooperations and combinations the first efficient, self-replicating prokaryote evolved on our planet Summary of the prebiotic broth & Organic chemical evolution theory 1. Formation of complex organic molecules, such as amino acids, nucleotides, occurs from simple precursor molecules, e.g. CO, H2, N2 in an early, oxygen-free Earth atmosphere and accumulate to form a prebiotic broth 2. Formation of the first complex polymers, e.g. nucleic acids, polypeptides, from organic monomers by dehydration synthesis in hot springs on charged surfaces, such as metal-containing rocks or clay formations 3. Establishment of the first primitive metabolic cycles on iron-nickel sulfides minerals 4. RNA established itself as the first self-replicating genetic material; in this “RNA world”, short strands of RNA copy themself accelerated with the help of Ribozymes 5. The first molecular cooperations (“coops”) form between many polymers and many other complex organic molecules 6. the first “useful” polypeptide chains show enzyme activity and trigger the evolution of reciprocal molecular cooperation between nucleic acids and proteins; RNA/protein coops form 7. formation of coops between self-replicating RNA/protein complexes and microscopic spheres enter the scene; so-called micelles made of early phospholipids create the first pre-biotic “cells” (= chambers) 8. Ongoing molecular evolution creates the first membrane-surrounded and selfreplicating primitive proto cells 14 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. 2. The pressurized iron-sulfur world theory this theory suggests a fast origin of life via a primitive autotrophic metabolism of small carbon-based molecules, such as pyruvic acid, acetic acid and alanine, in the hot and high pressure environment of deep ocean vents with the catalytic help of iron-nickel sulfur mineral surfaces (see section above) this idea is supported by a series of lab experimental evidence (see Corvis & Waechtershaeuser in the sections above) and the interesting observation that transition metal (e.g. nickel, iron, cobalt) sulfide clusters play a crucial role for the catalytic activity of many metabolically important enzymes in diverse groups of biological organisms also, iron-nickel-containing mineral sulfides are very common in deep ocean hydrothermal and volcanic vents 3. The panspermia (or cosmic ancestry) theory this theory favors the idea that life originated out in space and was carried to Earth in form of “seeds”/”endospores” traveling on/inside a meteorite; a meteorite impact early in Earth history may have triggered (“seeded”) the evolution of life this idea which recently gained increased popularity in the scientific community is supported by the discovery of extraterrestrial (ET) organic compounds in meteorites 1. the so-called Allende Meteorite, a meteorite which landed in 1969 near Allende, Mexico, was found to contain amino acids in its structure 2. a meteorite believed to be from Mars, discovered in 1996 in Antartica, shows several ET life-supporting features: it contains hydrocarbon molecules (the same as breakdown products of dead micro-organisms on Earth) its mineral phases are consistent with by-products of bacterial activity it shows tiny carbonate globules which may be microfossils of the primitive bacteria these findings also re-fueled the discussion about the possible existence of life elsewhere in the Solar System and the universe it also created further interest in the newly inaugurated biological sub-discipline of Astrobiology Future studies and discoveries will have to bring final proof which of the introduced theories turns out to be correct and best describes the currently enigmatic scenario which led to the development of life on our planet despite of the (currently) fragmented knowledge of the origin of life on our planet and the controversial discussions regarding the evolution of life, the earliest, traceable organisms evidently lived in form of primitive prokaryotes in the early oceans evidence of the first fossilized prokaryotes can be found in form of unique Precambrian rock formations called stromatolites, some of which have been dated back to more than 3 billion years 15 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. between 3.5 and 2.5 billion years, prokaryotes diversified already and appear in different shapes and forms in geological formations due to the metabolic activity of the photosynthesis-active cyanobacteria, the concentration of molecular oxygen (O2) increases in the Earth’s atmosphere this “first mass pollution” of the atmosphere in earth history led to a radical change in the bio-diversity on planet earth the increased oxygen concentration in the air triggered the mass extinction of anaerobic forms of life Under this slow but steady selective pressure and as a consequence, new, oxygen-adapted forms of life arise about 1.7 billion years ago the first ‘respirating’ eukaryotic cells appear on Earth and start their victorious evolution between 600 and 500 million years ago during the Cambrian period (590 – 505 mya), the first, primitive but structurally distinctive multi-cellular organisms appear which leave their traces in fossil records (see Images below) the first animal-like fossils appear at the end of the Proterozoic age (650 – 544 mya), also known as the Vendian period the first diversification of soft-bodied organisms, such as Dickinsonia, Tribrachidium and Leanchoilia illecebrosus, collectively known as the “Vendian fauna” or “Ediacaran fauna“ falls into this time period in many cases, scientists are not sure whether these ancient life forms were algae, lichens, giant protozoans, or even a separate kingdom of life Life form of the Cambrian period (590 – 505 million years ago) Leanchoilia illecebrosus 16 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. Life forms of the Cambrian period (590 – 505 million years ago) Image of fossil record Name Dickinsonia - Phylum ?? Tribrachidium - Cnidarian? - Echinoderm? about 500 – 400 million years ago the first primitive plants enter the scene in form of colonial green algae As mentioned at the very beginning of this chapter, today, more than 2 million different life forms or species call this planet home and testify the great success story of the great phenomenon called life Biologists classified these life forms into three major domains which include the Eubacteria (monerans), the Archaea bacteria and the Eukaryotes (see Graphic below) and seven kingdoms of life According to a very recently published findings, which are based on a bio-informaticsbased complete genome (DNA) analysis of selected prokaryotic and eukaryotic genomes, some scientists introduce the idea of a “ring of life” rather than to continue to speak of a tree of life with individual unconnected branches 17 SAN DIEGO MESA COLLEGE SCHOOL OF MATHEMATICS & NATURAL SCIENCES Intro Cell & Molecular Biology (BIOL 210); Instructor: Elmar Schmid, Ph.D. - - this new model showing the different major domains of life arranged in a connected ring considers the widely observed genetic phenomenon of horizontal gene transfer (“gene grapping”) amongst different species the ring includes following six major domains of life: 1. 2. 3. 4. 5. 6. Bacilli (Eubacteria) Cyanobacteria Proteobacteria Archaea bacteria Eocyta Eukaryota The 3 major domains of life on Earth 18