sábado, 13 de enero de 2007

A brief History of Life (10th grade assignment)

A Brief History of Life

Biologists studying evolution do a variety of things: population geneticists study the process as it is occurring; systematists seek to determine relationships between species and paleontologists seek to uncover details of the unfolding of life in the past. Discerning these details is often difficult, but hypotheses can be made and tested as new evidence comes to light. This section should be viewed as the best hypothesis scientists have as to the history of the planet. The material here ranges from some issues that are fairly certain to some topics that are nothing more than informed speculation. For some points there are opposing hypotheses -- I have tried to compile a consensus picture. In general, the more remote the time, the more likely the story is incomplete or in error.

Life evolved in the sea. It stayed there for the majority of the history of earth.
The first replicating molecules were most likely RNA. RNA is a nucleic acid similar to DNA. In laboratory studies it has been shown that some RNA sequences have catalytic capabilities. Most importantly, certain RNA sequences act as polymerases -- enzymes that form strands of RNA from its monomers. This process of self replication is the crucial step in the formation of life. This is called the RNA world hypothesis.

The common ancestor of all life probably used RNA as its genetic material. This ancestor gave rise to three major lineages of life. These are: the prokaryotes ("ordinary" bacteria), archaebacteria (thermophilic, methanogenic and halophilic bacteria) and eukaryotes. Eukaryotes include protists (single celled organisms like amoebas and diatoms and a few multicellular forms such as kelp), fungi (including mushrooms and yeast), plants and animals. Eukaryotes and archaebacteria are the two most closely related of the three. The process of translation (making protein from the instructions on a messenger RNA template) is similar in these lineages, but the organization of the genome and transcription (making messenger RNA from a DNA template) is very different in prokaryotes than in eukaryotes and archaebacteria. Scientists interpret this to mean that the common ancestor was RNA based; it gave rise to two lineages that independently formed a DNA genome and hence independently evolved mechanisms to transcribe DNA into RNA.

The first cells must have been anaerobic because there was no oxygen in the atmosphere. In addition, they were probably thermophilic ("heat-loving") and fermentative. Rocks as old as 3.5 billion years old have yielded prokaryotic fossils. Specifically, some rocks from Australia called the Warrawoona series give evidence of bacterial communities organized into structures called stromatolites. Fossils like these have subsequently been found all over the world. These mats of bacteria still form today in a few locales (for example, Shark Bay Australia). Bacteria are the only life forms found in the rocks for a long, long time --eukaryotes (protists) appear about 1.5 billion years ago and fungi-like things appear about 900 million years ago (0.9 billion years ago).
Photosynthesis evolved around 3.4 billion years ago. Photosynthesis is a process that allows organisms to harness sunlight to manufacture sugar from simpler precursors. The first photosystem to evolve, PSI, uses light to convert carbon dioxide (CO2) and hydrogen sulfide (H2S) to glucose. This process releases sulfur as a waste product. About a billion years later, a second photosystem (PS) evolved, probably from a duplication of the first photosystem. Organisms with PSII use both photosystems in conjunction to convert carbon dioxide (CO2) and water (H2O) into glucose. This process releases oxygen as a waste product. Anoxygenic (or H2S) photosynthesis, using PSI, is seen in living purple and green bacteria. Oxygenic (or H2O) photosynthesis, using PSI and PSII, takes place in cyanobacteria. Cyanobacteria are closely related to and hence probably evolved from purple bacterial ancestors. Green bacteria are an outgroup. Since oxygenic bacteria are a lineage within a cluster of anoxygenic lineages, scientists infer that PSI evolved first. This also corroborates with geological evidence.
Green plants and algae also use both photosystems. In these organisms, photosynthesis occurs in organelles (membrane bound structures within the cell) called chloroplasts. These organelles originated as free living bacteria related to the cyanobacteria that were engulfed by ur-eukaryotes and eventually entered into an endosymbiotic relationship. This endosymbiotic theory of eukaryotic organelles was championed by Lynn Margulis. Originally controversial, this theory is now accepted. One key line of evidence in support of this idea came when the DNA inside chloroplasts was sequenced -- the gene sequences were more similar to free-living cyanobacteria sequences than to sequences from the plants the chloroplasts resided in.
After the advent of photosystem II, oxygen levels increased. Dissolved oxygen in the oceans increased as well as atmospheric oxygen. This is sometimes called the oxygen holocaust. Oxygen is a very good electron acceptor and can be very damaging to living organisms. Many bacteria are anaerobic and die almost immediately in the presence of oxygen. Other organisms, like animals, have special ways to avoid cellular damage due to this element (and in fact require it to live.) Initially, when oxygen began building up in the environment, it was neutralized by materials already present. Iron, which existed in high concentrations in the sea was oxidized and precipitated. Evidence of this can be seen in banded iron formations from this time, layers of iron deposited on the sea floor. As one geologist put it, "the world rusted." Eventually, it grew to high enough concentrations to be dangerous to living things. In response, many species went extinct, some continued (and still continue) to thrive in anaerobic microenvironments and several lineages independently evolved oxygen respiration.

The purple bacteria evolved oxygen respiration by reversing the flow of molecules through their carbon fixing pathways and modifying their electron transport chains. Purple bacteria also enabled the eukaryotic lineage to become aerobic. Eukaryotic cells have membrane bound organelles called mitochondria that take care of respiration for the cell. These are endosymbionts like chloroplasts. Mitochondria formed this symbiotic relationship very early in eukaryotic history, all but a few groups of eukaryotic cells have mitochondria. Later, a few lineages picked up chloroplasts. Chloroplasts have multiple origins. Red algae picked up ur-chloroplasts from the cyanobacterial lineage. Green algae, the group plants evolved from, picked up different urchloroplasts from a prochlorophyte, a lineage closely related to cyanobacteria.
Animals start appearing prior to the Cambrian, about 600 million years ago. The first animals dating from just before the Cambrian were found in rocks near Adelaide, Australia. They are called the Ediacarian fauna and have subsequently been found in other locales as well. It is unclear if these forms have any surviving descendants. Some look a bit like Cnidarians (jellyfish, sea anemones and the like); others resemble annelids (earthworms). All the phyla (the second highest taxonomic category) of animals appeared around the Cambrian. The Cambrian 'explosion' may have been a result of higher oxygen concentrations enabling larger organisms with higher metabolisms to evolve. Or it might be due to the spreading of shallow seas at that time providing a variety of new niches. In any case, the radiation produced a wide variety of animals.

Some paleontologists think more animal phyla were present then than now. The animals of the Burgess shale are an example of Cambrian animal fossils. These fossils, from Canada, show a bizarre array of creatures, some which appear to have unique body plans unlike those seen in any living animals.

The extent of the Cambrian explosion is often overstated. Although quick, the Cambrian explosion is not instantaneous in geologic time. Also, there is evidence of animal life prior to the Cambrian. In addition, although all the phyla of animals came into being, these were not the modern forms we see today. Our own phylum (which we share with other mammals, reptiles, birds, amphibians and fish) was represented by a small, sliver-like thing called Pikaia. Plants were not yet present. Photosynthetic protists and algae were the bottom of the food chain. Following the Cambrian, the number of marine families leveled off at a little less than 200.
The Ordovician explosion, around 500 million years ago, followed. This 'explosion', larger than the Cambrian, introduced numerous families of the Paleozoic fauna (including crinoids, articulate brachiopods, cephalopods and corals). The Cambrian fauna, (trilobites, inarticulate brachiopods, etc.) declined slowly during this time. By the end of the Ordovician, the Cambrian fauna had mostly given way to the Paleozoic fauna and the number of marine families was just over 400. It stayed at this level until the end of the Permian period.

Plants evolved from ancient green algae over 400 million years ago. Both groups use chlorophyll a and b as photosynthetic pigments. In addition, plants and green algae are the only groups to store starch in their chloroplasts. Plants and fungi (in symbiosis) invaded the land about 400 million years ago. The first plants were moss-like and required moist environments to survive. Later, evolutionary developments such as a waxy cuticle allowed some plants to exploit more inland environments. Still mosses lack true vascular tissue to transport fluids and nutrients. This limits their size since these must diffuse through the plant. Vascular plants evolved from mosses. The first vascular land plant known is Cooksonia, a spiky, branching, leafless structure. At the same time, or shortly thereafter, arthropods followed plants onto the land. The first land animals known are myriapods -- centipedes and millipedes.

Vertebrates moved onto the land by the Devonian period, about 380 million years ago. Ichthyostega, an amphibian, is the among the first known land vertebrates. It was found in Greenland and was derived from lobe-finned fishes called Rhipidistians. Amphibians gave rise to reptiles. Reptiles had evolved scales to decrease water loss and a shelled egg permitting young to be hatched on land. Among the earliest well preserved reptiles is Hylonomus, from rocks in Nova Scotia.

The Permian extinction was the largest extinction in history. It happened about 250 million years ago. The last of the Cambrian Fauna went extinct. The Paleozoic fauna took a nose dive from about 300 families to about 50. It is estimated that 96% of all species (50% of all Families) met their end. Following this event, the Modern fauna, which had been slowly expanding since the Ordovician, took over.

The Modern fauna includes fish, bivalves, gastropods and crabs. These were barely affected by the Permian extinction. The Modern fauna subsequently increased to over 600 marine families at present. The Paleozoic fauna held steady at about 100 families. A second extinction event shortly following the Permian kept animal diversity low for awhile.
During the Carboniferous (the period just prior to the Permian) and in the Permian the landscape was dominated by ferns and their relatives. After the Permian extinction, gymnosperms (ex. pines) became more abundant. Gymnosperms had evolved seeds, from seedless fern ancestors, which helped their ability to disperse. Gymnosperms also evolved pollen, encased sperm which allowed for more outcrossing.

Dinosaurs evolved from archosaur reptiles, their closest living relatives are crocodiles. One modification that may have been a key to their success was the evolution of an upright stance. Amphibians and reptiles have a splayed stance and walk with an undulating pattern because their limbs are modified from fins. Their gait is modified from the swimming movement of fish. Splay stanced animals cannot sustain continued locomotion because they cannot breathe while they move; their undulating movement compresses their chest cavity. Thus, they must stop every few steps and breath before continuing on their way. Dinosaurs evolved an upright stance similar to the upright stance mammals independently evolved. This allowed for continual locomotion. In addition, dinosaurs evolved to be warm-blooded. Warmbloodedness allows an increase in the vigor of movements in erect organisms. Splay stanced organisms would probably not benefit from warm- bloodedness. Birds evolved from sauriscian dinosaurs. Cladistically, birds are dinosaurs. The transitional fossil Archaeopteryx has a mixture of reptilian and avian features.

Angiosperms evolved from gymnosperms, their closest relatives are Gnetae. Two key adaptations allowed them to displace gymnosperms as the dominant fauna -- fruits and flowers. Fruits (modified plant ovaries) allow for animal-based seed dispersal and deposition with plenty of fertilizer. Flowers evolved to facilitate animal, especially insect, based pollen dispersal. Petals are modified leaves. Angiosperms currently dominate the flora of the world -- over three fourths of all living plants are angiosperms.
Insects evolved from primitive segmented arthropods. The mouth parts of insects are modified legs. Insects are closely related to annelids. Insects dominate the fauna of the world. Over half of all named species are insects. One third of this number are beetles.

The end of the Cretaceous, about 65 million years ago, is marked by a minor mass extinction. This extinction marked the demise of all the lineages of dinosaurs save the birds. Up to this point mammals were confined to nocturnal, insectivorous niches. Once the dinosaurs were out of the picture, they diversified. Morgonucudon , a contemporary of dinosaurs, is an example of one of the first mammals. Mammals evolved from therapsid reptiles. The finback reptile Diametrodon is an example of a therapsid. One of the most successful lineages of mammals is, of course, humans. Humans are neotenous apes. Neoteny is a process which leads to an organism reaching reproductive capacity in its juvenile form. The primary line of evidence for this is the similarities between young apes and adult humans. Louis Bolk compiled a list of 25 features shared between adult humans and juvenile apes, including facial morphology, high relative brain weight, absence of brow ridges and cranial crests.

The earth has been in a state of flux for 4 billion years. Across this time, the abundance of different lineages varies wildly. New lineages evolve and radiate out across the face of the planet, pushing older lineages to extinction, or relictual existence in protected refugia or suitable microhabitats. Organisms modify their environments. This can be disastrous, as in the case of the oxygen holocaust. However, environmental modification can be the impetus for further evolutionary change. Overall, diversity has increased since the beginning of life. This increase is, however, interrupted numerous times by mass extinctions. Diversity appears to have hit an all-time high just prior to the appearance of humans. As the human population has increased, biological diversity has decreased at an ever-increasing pace. The correlation is probably causal.

Q&A.

1. List and describe 10 areas of study in Biology. (Ex: evolutionary biology)

2. Describe the celluar differences between the prokaryotes ("ordinary" bacteria), archaebacteria (thermophilic, methanogenic and halophilic bacteria) and eukaryotes. Eukaryotes include protists (single celled organisms like amoebas and others).

3. Where do scientists believe life originated?

4. Modern biology is based on several great ideas, or theories: The Cell Theory, Theory of Evolution by Natural Selection, Gene Theory, Homeostasis. Write the main statements that each of these theories hold.

5. Explain the importance of photosynthesis in nature.

6. What events happened during the “Cambrian explosion”?

7. Why do scientists believe flowering plants evolved?

8. What are endosymbionts? What organelles in modern cells are believed to have evolved from endosymbiontic organisms?

9. When did the first animals appear on Earth?

10. Contrast Creationism and Evolution. Write a short essay explaining both points of view; state your personal opinions (conclusions) and the reasons why you came to that conclusion.

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