Transcript Document
A brief history of life Peter Shaw We are lucky enough to emerge from the most remarkable lump of stone in the universe (at least as far a we know). We know that the universe exploded from a Big Bang about 14 billion years ago, and is widely if thinly seeded with hydrogen, helium and lumps of stone. As far as we know, all the ones we have studied have their surfaces in simple accord with basic physical chemistry. The surfaces of the moon, and Mars, contain simple inorganic chemicals in utter equilibrium with their surroundings. Earth is different. The air is not only thick (thanks to earth’s magnetic field), but an unstable mix of N2 and O2. These should end up as nitrate in the sea. Some bits of the earth’s surface burn given half a chance! The sea is oddly deficient in some ions, notably ammonium, nitrate and potassium compared to chemical equilibria. All these are the signatures of Life. Biological life, filtering air and water into bodies, and cycling bodies through food chains into soils. Today I want to give you a brief history of this most remarkable transformation. We rely only on evidence: fossils to tell us what conditions were like, extant life forms to infer common ancestry, chemical simulations to explore what might have happened. Sadly these will probably never ell us the whole story. The most important sections concern details of biochemical organisation in proto-life, and we simply do not know, sorry. How old? We can get reasonably reliable dates commensurate with the planet’s age using very slow radioactive decay processes. The best is the decay of K40 to Ar40, which has a half life around 1.270,000,000 years (1.27E9 years). Ar is a highly inert gas so gases off – unless locked inside a glassy volcanic stone. Thus the Ar40:K40 ratio of volcanic stones is an excellent clock for when the lava cooled. The oldest stones on the planet – tiny grains of the gem zircon inside sandstone formed in an 4BYBP ocean (probably before life) give dates around 4.3BYBP. Note that this is about 1/3 age of the universe. Current thinking suggests true earth age around 4.5 BYBP. Age of your planet: 4,500,000,000 years. Hadean Eon Earth assembled from plantary rubble, and experienced a partial melt when something (theia) bashed into us to knock off our moon. Models suggest that this helped dense iron-rich materials sink to the core, giving us a magnetic field. Oceans are projected to have started to form c. 4.4 BYBP, with the gradual settling down of a surface crust. With little atmosphere and intense instability we assume earth’s surface was lifeless for most of this phase. In the beginning Then something remarkable happened. We had waterbased seas with organic compounds deposited from comets, CO2 + CH4 in the atmosphere, irradiated with UV. We know that in the lab simple building blocks of life appear - sugars, amino acids. Somewhere in this mix, a chemical reaction occurred which ultimately promoted its own copying. Nowadays these copying reactions centre on DNA; early versions may well have relied on RNA, earlier still maybe clay crystals. It is quite possible that the reactions involved first appeared on Mars, which later seeded us with a spalledoff meteor. Somehow, an early reaction took ambient energy to copy itself. LUCA We can only go on evidence, and the evidence is that modern-looking filamentous prokaryotes (loosely; algae) appeared very early on, within 700,000,000 years of earth’s formation. We do not know about their biochemistry, nor about the intricate dance of chemical reactions that preceded them. The evidence of modern life forms is clear; all extant life forms came from one ancestral design. LUCA - the Last Universal Common Ancestor. One system bundled itself into phospholipid bubbles and coded itself on DNA with an arbitrary mapping that has permeated all life. A question that repeatedly gets thrown at biologists is something along the lines of “What is the purpose of life?” (Often by people who insist on disbelieving evolution..) Isolate out the dual meaning of ‘life’ - your personal goals are your personal choice and irrelevant to evolutionary theory - and the unask the question. There is no evidence to suggest that the set of self-copying chemical reactions we call ‘Life’ has or ever had a purpose. Purpose implies planning, a conscious overview, which is why it is so important to deny theists the use of the term. Analogies Take a great pile of randomly assorted pebbles and dump them on the edge of the sea, somewhere with wave action. Their size distribution will be random. 3 scenarios now; 1: Too little energy is applied; the stones remain unsorted. No pattern persists. 2: Too much wave energy destroys the system, here by removing stones. No patterns persist. 3: The wave energy is enough to move the stones around. A pattern appears - big stones at the top, smallest down the base. This is the usual pattern observed on most stony beaches by the sea. What is the purpose of that pattern? Carbon chemistry Take a sterile planet with organic molecules in liquid water in its oceans, and apply UV radiation capable of ionizing molecules. 3 scenarios now; 1: Too little energy is applied; no interesting reactions take place, no new chemistry appears. No pattern visible. 2: Too much energy destroys the system, here by destroying molecules or boiling the water. No patterns persist. 3: The solar energy is just enough to allow a self-copying reaction based on solar-derived chemical changes. A pattern appears - proto life. What is the purpose of that pattern? Archean and proterozoic Eras This period – the majority of earth’s history – ran from 3.8-0.7 BYBP. Only stromatolites, no signs of animals. The fossil record tells us that the majority of earth’s history it supported oceans with slimy algae forming dirt-trapping hummocks. About 3 Billion years, at least 20* the rule of the dinosaurs. Just stromatolite fossils, and slowly increasing oxygen levels. Stromatolites These are our oldest fossils but can still be found (notably Shark Bay in Australia). Basically they are mounds made by photosynthetic algae which entrap sediment so accrete, and can become quite large (a metre or so) in shallow water. Humble, but we owe our existence to their slow diligent production of oxygen. About 3800 million years ago… The earth had oceans, life, but the air was CO2/CH4 without oxygen. The oceans contained vast amounts of dissolved Iron II (blue-back). The earliest fossils are of stromatolites (+ strange planktonic remains called acritarchs). Age of planet earth, billions of years 4.5 4 3.5 3 2. 5 2 1.5 1 Photosynthesis Oxygen builds up in air, removing Iron 2 from oceans as rust and filtering UV 0.5 0 Animals Dinosaurs Endosymbiosis One more really important thing happened sometime in the proterozoic, and we don’t know when. One cell swallowed another, but instead of digesting it as dinner, the intruder took up permanent residence using up a toxic waste gas oxygen and creating high energy compounds. This gave us the eukaryotic cell, with a nucleus and organelles. All later multicellular life came from one ancestral eukaryotic fusion. This process has happened again, notably when a eukaryotic cell ingested a cyanophyta cell and kept it to evolve into chloroplasts - the light capturing organs of all higher plants. Endosymbiosis For many years people knew that chloroplasts looked like some bluegreen algae (cyanophyta, which are prokaryotes) down to the organisation of their stroma. But they weren’t invading pathogens, they are well defined cellular organelles, copied faithfully down the generations. Oddly, another well-defined organelle, the mitochondrion, also looks a bit like a bacterial cell. Odd that, but just one of those oddities… Then in the 1960s Lynn Margulis produced her thesis suggesting that these organelles really are prokaryotes, living as endosymbionts inside their host cell. (And was laughed at for about 20 years..) Endo – inside Symbiosis – a mutually beneficial relationship between two life forms. Lynn Margulis predicted that these autonomous organelles would have independent DNA, and to widespread amazement by the early 1980s it became clear that chloroplasts and mitochondria DID have their own DNA, in a circular loop just like a prokaryote. The endosymbiosis heresy became established fact. (This is where science differs from religion.) They also have their own ribosome. Specifically, chloroplasts can be visualised as a free-living cyanophyta which has been swallowed (invaginated) by a host cell but remained undigested. Large cell, perhaps of an archaean Photosynthetic cyanophyta Photosynthesis continues, inside a vacuole and inside its own cell wall. A Mitochondrion (the tubby-sausage silhouette is typical). Note the dual membrane system, with the inner membrane equating to the ancestral cell wall of the endosymbiotic bacteria. Chloroplasts Again note the double membrane, the inner a relic of the prokaryotic endosymbiont. 4-10μm Bodies! The buildup of O2 had several profound consequences. 1: It protected from UV, enabling life to emerge from under deep water. 2: It enabled the rapid metabolisms of multi-cellular organisms. Ediacaran or Vendian era First fossils appearing to animal are the bizarre Ediacaran creatures that briefly flourished 700-450 MYBP. They only persist as enigmatic fossils, fundamentally unlike anything since. The strange name comes from the Ediacara range in S Australia where the definitive faunas come from. The first however was found by a schoolboy in Charnwood forest (UK), at a time when everyone knew that fossils in precambrian deposits were impossible. The genus is called Charnia to this day. We know nothing about these animals? The basal bulb probably anchored them in a mat of microbial filaments? The Cambrian explosion The fossil evidence of animals first appears with shelly forms at the dawn of the Cambrian, c. 540-490 MYBP. We know from a truly remarkable deposit called the Burgess Shales that diverse fauna, justabout recognisable as modern forms in most cases, existed about 500 MYBP. The key point is the appearance of hard shells, allowing good fossilisation, but we have enough Vendian fossils by now to be confident that there were few if any hard-bodied anythings then. Age of planet earth, billions of years 4.5 4 3.5 3 2. 5 2 1.5 1 Photosynthesis Oxygen builds up in air, removing Iron 2 from oceans as rust and filtering UV 0.5 0 Animals Precambrian Aysheaia – our ancestor Burgess shales fauna Hallucinigenia In the early days of animals the seas held tadpole-like protofish, shrimp-like arthropods, jellyfish corals etc, molluscs (and lamp shells, not molluscs at all). Probably lots of algae, hard to know. On land: nothing. The devonian was the age of fish - the most advanced life forms on the planet were armoured fish, co-predating with giant sea-scorpions. Corals though present were often superceded by sea lilies (crinoids). Land colonisation Our 1st land fossils come around 390MYBP, notably the amazing rhynie cherts of Scotland, showing us the cells and stomata of plants before leaves evolved, and how little springtails have changed. Asteroxylon mackei, an early lycopod. X = xylem, t = leaf traces, scale bar 1mm A modern club moss Lycopodium for comparison. They also have striking xylem (see slide). Geological periods The recent (last 700 MYBP) history of the planet is named by geologists, but using biologists’ data. Geological eras are defined by their fossil assemblages, which is observed to under occasional massive abrupt changes. These are Mass extinction events - MEEs, and they set boundaries; silurian, devonian, carboniferous, permian, triassic, jurassic, cretaceous, palaeocene, eocene, .. holocene. MEE – mass extinction events. Punctuation marks in the earth’s history. We tell of slow evolutionary processes and geological time scales, just as we talk of societies advancing gradually towards [happiness, democracy, equality…] Then recall Baghdad May 2003; no society, no structures, collapse and meltdown. MEEs are when the biosphere does the same thing. Geological Boundaries Geologists divide the planet’s history up into bite-sized divisions. These are NOT merely to satisfy vanity – “Let’s make up a name for the period from 400-300 MYBP”. These reflect meaningful differences in the stones based on their fossil content. If you see a trilobite in the shales the rock’s age lies between two clear boundaries. No hard-bodied animals before the Cambrian, no Trilobites after the Permian. The names were largely coined before isotopic dating had been developed, and latterly there was huge tension between biologists (who needed long timescales) and physical geologists (who calculated how long the earth would take to cool from sun surface temperatures; c. 20 million years) Geological time scales coded as: Era: Period: Epoch Precambrian: = most of history + Ediacaran (Vendian), 4500570 MYBP Paleozoic: 570-225 MYBP; Cambrian Ordivician Silurian Mesozoic: Cenozoic: Precambrian Devonian Carboniferous Permian Triassic Cretaceous Jurassic Cretaceous Tertiary (Palaeocene, Eocene, Oligocene Miocene Pliocene) Quaternary (Pleistocene holocene) Palaeozoic Mesozoic Cenozoic shelled animals 700 600 500 Coal 400 300 Age MYBP ? Anthropocene Oil 200 100 0=now Geological boundaries As explained, are marked by indicator fossils. These can be whatever is fossilised well and often, invariably hard bodied forms: These include a roll call of taxa that were world-dominating in their time, and have now vanished, either absolutely or in ecosystem dominance: Trilobites, Conodonts, ammonites – extinct Graptolites, sea lilies, brachiopods – survive at a low level. The Permian MEE The Palaeocene / Mesozoic boundary occurs between the Permian and the Triassic, 251 MYBP. It marks the greatest mass killing in the Earth’s history, with >95% of known marine species vanishing at the boundary. The most famous were the Trilobites. (Also 70% terrestrial spp, including gorgonopsian reptiles) Sediments around the boundary show signs of anaerobiosis, and there is a strong suggestion that oxygen tensions fell so low that life became impossible for many life forms. We don’t really know why, but the evidence below points one way. There is no meteor crater whose date fits, little chemical evidence from the boundary suggesting asteroid material (some contested argon ratios in 2 sites), but definite synchronous large volcanic eruptions (forming the Siberian traps). CO2 trends over time RCO2 (1* = 350 ppm) 0 5 10 15 20 25 = methane catastrophe P/T 600 500 400 Toarcian 183mybp PETM 55mybp 300 200 100 0=now Age MYBP Palaeozoic Mesozoic Cenozoic The K/T boundary The mesozoic / cenozoic boundary is marked by the K/T boundary (= Keuper / Tertiary). This is dated to 64MYBP, and is marked by a faunal shift nearly as great as the Permian. The boundary is observed around the world as a homogenous ashy deposit, a few cm – 10s cms deep. This is in itself unique: there are NO OTHER global boundaries that manifest as a horizon over the whole planet. Below the layer are mesozoic reptiles. In Hell Creek you can find bones right up to the layer (some claim density falls away near the boundary, but this looks like a sampling artefact; bones that big take a long time to get covered over.) Above the ashy layer, no mesozoic reptiles, and not much else: a spike of ferns, some insects, then mammals start to radiate. For this event we know that a major extraterrestrial impact was culpable, in part. There is a chemical signal, detected and identified by the Walter and Luis Alvarez (father and son). It concerns the element iridium, a siderophilic mineral rarely found on earth but relatively enriched in asteroidal material. (I don’t know why BTW). Its isotopic signature matched an asteroid too. The KT boundary in Alberta badlands The K/T boundary turned out to be enriched in Iridium 10-15* background levels, all over the planet. This was convincing evidence that an asteroid had hit earth at the K/T boundary, leaving detritus in the ashy layer. The iridium values gave the asteroid a 10km diameter The unifying theme Time and again after a MEE there is a lull while new species evolve, then the system settles down to land plants supporting large herbivorous land animals, that in turn support predators (and presumably scavengers and parasites). This didn’t apply in the carboniferous, but has done every time since. Spot the modern parallels: Permian 290-250 MYBP Land plants herbivores carnivores dicynodonts Gorgonopsids Seed ferns (here Glossopteris) (note sabre teeth) Jurassic-Cretaceous 210-64 MYBP Land plants Gingo, cycads herbivores sauropods carnivores Tyranosaurus rex Recent food chain (where?) Land plants grass herbivores litopterns carnivores Sabre tooth marsupial tiger Thylacosmilus (replaced by Smilodon) Life in an MEE We are probably living in the early stages of a MEE to rival the K/T boundary. The future biosphere + climate system will be the anthropocene, dominated by an intense heat pulse (CO2 -> CH4 release). Recovery from an MEE takes c. 10 million years. Remember that when telling your kids that you recall the Chinese river dolphin being found to be extinct. Lipotes vexillifer Extinct 2006