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Chapter 29: Early Evolution and the Procaryotes


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Chapter 29: Early Evolution And The Procaryotes

A. THE ORIGIN OF BIOLOGICAL SYSTEMS

29.1 Elements are formed through the natural evolution of stars.

a. According to contemporary cosmology, the universe has been undergoing a cosmic evolution since its origin about 13 billion years ago.

b. Hydrogen and helium nuclei form rocky planets like earth and are also the substance of the organic molecules of which organisms are made.

c. A planet must maintain just the right range of temperatures to assure that water remains a liquid on the surface.

d. One might speculate that a galaxy like ours, the Milky Way, must hold vast numbers of planets that are suitable for life.

29.2 Organic molecules form in the reducing atmosphere of primitive planets.

a. In the 1930s, A. I. Oparin, developing an argument that was also presented by J. B. S. Haldane in 1929, pointed out that the primitive earth must have had a reducing–rather than an oxidizing–atmosphere, made mostly of hydrogen, methane, ammonia, nitrogen, and water.

b. In 1953, Stanley Miller tested the Haldane—Oparin hypothesis using the apparatus shown in Figure 29.1.

1. Miller tried to reconstruct primitive conditions by passing electrical sparks through a mixture similar to the hypothetical primitive atmosphere.

2. Miller found that after several days, the mixture in the apparatus had formed quite a variety of organic compounds, including the common amino acids.

3. Miller's experiment has been repeated many times with various initial mixtures, always with similar results.

c. Sidney Fox found that simply heating dry mixtures of amino acids produces so-called "proteinoids," small proteins (3—10 kDa) with many properties of modern proteins, including some ability to catalyze chemical reactions.

1. Fox and his colleagues later found that hot, salty solutions of peptides form microspheres as they cool.

2. Microspheres are stable nonliving structures with many properties of modern procaryotic cells and about the same size (about 2 µm in diameter).

d. The formation of cells must occur early in evolution, for two reasons.

1. Further evolution requires that metabolism be confined to small, enclosed spaces where high concentrations of metabolites and enzymes can develop.

2. Genetic systems must be confined to cells to take advantage of mutational novelties; evolution depends on competition among individuals (cells or organisms), and this could not happen if each genetic novelty were shared with a large system.

 

29.3 The evolution of a functioning genome is problematic.

a. The most critical feature of an organism is its genome since evolution depends on the selection of individuals with variant genomes.

b. A genome must replicate and direct the synthesis of proteins.

c. Although modern cellular genomes are all DNA, the first nucleic acids to become functional were probably RNA.

1. RNA molecules have an inherent ability to interact with one another through base-pairing, so they can replicate as DNA does.

2. RNA molecules can also fold into complicated forms and act as ribozymes which are RNA molecules that catalyze chemical reactions just as protein enzymes do.

3. During the RNA-world stage of evolution, many of the functions now performed by proteins must have been performed by RNA molecules.

d. Francis Crick and Leslie Orgel have suggested that two kinds of RNA might have evolved early on (Figure 29.2).

1. Some RNAs had the potential to become genomes ("protogenomes") because their base sequences allowed them to replicate with particular ease.

2. Other RNAs, having an affinity for specific amino acids, then served as primitive transfer RNAs by helping to line up amino acids along a protogenome, thus catalyzing a simple protein synthesis.

29.4 We can reconstruct a likely course of procaryote evolution.

a. The earliest organisms must have been heterotrophs that fed on preformed organic compounds (Figure 29.3).

1. These cells must have evolved pathways for mobilizing energy through fermentation, probably including the glycolytic pathway, producing ATP by substrate-level phosphorylation, with no electron transport system.

2. Suppose, at this early stage of evolution, a critical monomer became depleted. Only the organisms that had an enzyme for converting another compound into the one that was depleted survived.

3. Through mutation and natural selection, pathways for biosynthesis evolved, and organisms gradually became more independent (Figure 29.4).

b. Photosynthesis evolved with the ability to make porphyrins, including chlorophyll.

1. Plant photosynthesis, which produces oxygen as a byproduct (oxygenic photosynthesis), is an advanced form that requires two photosystems and must have taken a long time to evolve.

2. Around 2.0—1.8 billion years ago, phototrophs probably used various forms of anoxygenic photosynthesis, which does not produce oxygen as a byproduct.

3. The first phototrophs probably used a form of photosystem I just to oxidize organic substrates. This text refers to these organisms as stage 1.

4. The modern survivors of that retain stage-1 photosynthesis are green sulfur bacteria and purple non-sulfur bacteria (Figure 29.5).

5. Stage-2 photosynthesis came about with the evolution of the PCR (Calvin—Benson) cycle.

6. Modern purple sulfur bacteria use stage-2 photosynthesis and can only grow anaerobically in the light using H2S as an electron donor instead of H2O (Figure 29.6).

7. With the development of photosystem II, oxygenic photosynthesis finally evolved (stage 3).

8. Stage-3 photosynthesis is used in modern cyanobacteria (blue-green bacteria; Figure 29.7).

9. Cyanobacteria have the same chlorophyll a as eucaryotic phototrophs, and they produce oxygen as a byproduct of photosynthesis.

c. The production of oxygen opened the door to aerobic respiration.

1. Oxygenic photosynthesis began to shift the atmosphere from reducing to oxidizing.

2. The evolution of cytochrome enzymes that could reduce oxygen to water was a great event that created modern aerobic respiration.

3. Since the DNA genome is vulnerable to destruction by oxygen, it was advantageous to sequester it in a nucleus and to confine oxygen utilization and production to separate organelles, to mitochondria and chloroplasts.

4. When cells evolved mitochondria and chloroplasts, they could become larger, with local regions of the cytoplasm served by these ATP factories.

 

B. THE KINGDOM MONERA

29.5 Procaryotic cells have no true nucleus and are usually very small.

a. The kingdom Monera includes all procaryotes except Archaebacteria (see Sidebar 29.1).

1. Most procaryotes are single cells with diameters on the order of 1 µm or less.

2. Some procaryotes are as large as 10—30 µm in diameter and one recently-discovered bacteria has cell lengths of about 300 µm (Figure 29.9).

b. Procaryotic chromosomes are compacted into nucleoids, not surrounded by a nuclear envelope, and they have no endoplasmic reticulum or mitochondria.

1. Each procaryotic cell bears a single chromosome, usually circular.

c. Procaryotic flagella, if present, are simple protein rods, not 9 + 2 complexes of microtubules.

d. Most procaryotes have a cell wall surrounding the plasma membrane.

1. Bacterial cell walls are made of a murein, or peptidoglycan, which is a polymer made by bonding two kinds of monomers into a two-dimensional network (Figure 29.10).

2. Mureins are the largest covalently bonded molecules in the world.

3. Bacterial cell walls are often rigid but porous, and they give bacteria their characteristic shapes.

29.6 Bacteria may be spheres, rods, spirals, or long filaments.

a. Much can be learned about bacteria morphology through their names.

1. Bacteria can be a general term used for all procaryotes, but it is not very descriptive.

2. Bacteria often exist in the form of rods, spirals (spirilla, singular spirillum) and curved cells, and spheres (cocci, singular coccus) (Figure 29.11).

3. The shapes of the bacteria often provide both the formal and informal names of bacteria such as diplococci (pairs of cocci), streptococci (chains of cocci), and staphylococci (random clumps of cocci).

b. Spirochetes are soft-walled, flexible procaryotes that zip along through water like tiny corkscrews (Figure 29.12).

1. Spirochetes propel themselves by contracting an axial filament made of fibers similar to bacterial flagella.

2. Some spirochetes are 500 µm long, yet they are thin enough (0.1—0.6 µm in diameter) to pass through filters that remove most bacteria.

3. Most spirochetes are soil and water organisms.

4. Some spirochetes are pathogens, such as Treponema pallidum, the causative agent of syphilis.

c. The slime bacteria or Myxobacteria are another group of procaryotes with flexible cell walls.

1. Slime bacteria move with a gliding mechanism and are common in soils, on rotting materials and in animal dung (Figure 29.13).

2. Myxobacteria aggregate into mushroom-like clusters, which sometimes become quite large.

3. Stalked bacteria (Figure 29.14) are common in contaminated water.

d. Mycobacteria include the agents that cause tuberculosis and leprosy.

1. Mycobacteria cells are irregular and branched with unusual cell walls containing complex lipids.

2. Actinomycetes, which are closely related to mycobacteria, form long, branched, filaments with many nucleoids, but they are coenocytic because they lack cross-walls or have only incomplete walls (Figure 29.15).

3. Streptomyces, a representative actinomycete, reproduces by forming spores, each containing a chromosome, at the end of each filament.

e. The procaryotes include mycoplasmas, the smallest of cells, with diameters of only 0.1—0.3 µm (Figure 29.16).

1. Mycoplasmas lack cell walls and can grow into irregular elongated forms.

2. Mycoplasmas contain the minimal apparatus needed for metabolism and reproduction.

3. Some mycoplasmas are harmless while others are the agents of plant and animal diseases such as pleuropneumonia, a lung infection of cattle.

29.7 Bacteria are classified by cell shape, metabolism, and reaction to the Gram stain.

a. Bergey's Manual of Determinative Microbiology is an excellent consensus on procaryotic taxonomy.

b. Table 29.1 illustrates the major groups of procaryotes.

c. The major group of procaryotes are defined primarily by three criteria.

1. The first criterion is cell morphology.

2. The second criterion is metabolic pattern (Figure 29.17).

3. The third criterion for classifying bacteria is their reaction to the Gram stain (Figures 29.19 and 29.20).

d. Organisms can be distinguished as obligate or facultative.

1. An organism's way of life is obligate if it is obliged to live a certain way with no other options.

2. An organism's way of life is facultative if it can live one or more different ways.

3. Obligate anaerobes, for instance, can only live in environments with no trace of oxygen, while facultative anaerobes can survive either with or without oxygen (Figure 29.18).

 

C. THE ECOLOGY AND USES OF BACTERIA

29.8 Bacteria have critical roles in every ecosystem.

a. Bacteria are most significant as decomposers in the ecosystem, organisms that reduce wastes to simpler materials.

1. Bacteria (and molds) produce extracellular hydrolytic enzymes that attack the polymers of these wastes, releasing monomers on which they, and others that share their living space, can grow.

2. Bacteria in turn become food for small animals, and thus some of their mass is recycled back into the food chain.

b. Bacteria that live on unusual transformations of inorganic compounds also play important roles in the cycling of materials in ecosystems, performing critical steps in the nitrogen and sulfur cycles.

1. Sometimes two or more species of bacteria cooperate fortuitously to degrade an unusual organic compound step-wise by combining their enzymatic capabilities, although each one by itself is unable to grow on the compound.

2. Figure 29.21 shows a simple cycle between a phototrophic bacterium and a sulfate-reducing bacterium.

3. Figure 29.22 shows a similar cycle between a chemoautotroph that reduces CO2 to organic compounds and a nitrogen fixer that produces nitrogenous compounds.

4. Through this kind of mutualism, the two organisms grow together much better than either could grow alone.

29.9 Many bacteria cause infectious diseases.

a. The etiology of disease–its underlying cause or causes–is either functional or infectious.

1. Functional disease, such as heart disease, results from a malfunction in the organism itself.

2. Infectious disease, is due to some other organism or virus that grows as a parasite, obtaining its nourishment on the surface of the host (ectoparasitism) or somewhere within its intestinal tract or other cavities, or even within its tissues (endoparasitism).

3. Diseases are caused by bacteria, protozoans, fungi, worms, and viruses.

4. Some small bacteria can even grow inside larger bacteria (Figure 29.23).

b. We owe our modern understanding of contagious diseases largely to the research of Louis Pasteur.

1. In 1835, Charles Cagniard-Latour and Theodor Schwann showed independently that fermentation in beer and wine is caused by the growth of a microorganism, yeast.

2. Chemists of the day found the Cagniard-Latour/Schwann findings to be totally ridiculous. The chemists believed that fermentation was a purely chemical process.

3. Pasteur showed that by transferring yeasts and bacteria from one culture to another, would transfer the fermentative activity.

4. Pasteur also showed that when wines become sour it is because they are contaminated with undesirable bacteria.

5. In order to counter such contamination, Pasteur invented the process now called pasteurization, in which the wine (or other food such as milk) is heated briefly, just enough to kill the responsible organisms while leaving the food unharmed.

c. Pasteur realized that animal diseases might also result from infection by microorganisms.

1. In the 1860s, he demonstrated the infectious agents were indeed responsible for silkworm diseases, which at the time were ravaging the French silk industry.

2. In the 1870s, Pasteur and Robert Koch showed that bacteria were causing anthrax, a disease that can strike many mammals, including humans.

3. Koch found that anthrax bacteria form protective spores that may endure for a long time, so animals may be infected merely by grazing in certain pastures.

d. Koch devised a set of four conditions, now known as Koch's Postulates, that must be satisfied before a disease can be positively attributed to some organism.

1. The organism must be recovered from animals that have the disease.

2. The organism must be grown in pure culture.

3. When healthy animals are inoculated with this culture, they must contract the disease.

4. The organism must again be recoverable from an inoculated animal.

e. Using Koch's Postulates, the etiologic agents of many diseases were identified during the next decades.

f. Around 1864, the English surgeon Joseph Lister realized that infectious agents might be responsible for the high rate of sepsis (decay and death of tissues) associated with surgery, and he instituted procedures such as the sterilization of surgical instruments and the use of disinfectants.

29.10 Pathogens produce disease through invasion and toxin production.

a. Pathogens are either invasive, causing disease by invading the host and growing in its tissues, or toxigenic, causing disease by producing toxins, or poisons.

1. Clostridium perfringens is an extremely invasive pathogen that causes gas gangrene. Clostridia form spores that may lie dormant in the soil for a long time until they are carried into a wound, especially a deep puncture wound. If untreated, the cell- and tissue-destroying bacteria thrives leaving decaying tissue behind.

2. Clostridium botulinum, the agent of botulism, produces the botulinus toxin that paralyzes the neuromuscular junctions where nerve endings contact muscles.

29.11 Infectious agents are transmitted to new hosts from reservoirs of infection.

a. A growth of parasitic organisms in some larger host, with the accompanying destruction of tissues and production of toxic byproducts, produces an infection.

b. Every surface of a multicellular organism is a specialized ecological niche where something else can grow.

1. Most organisms achieve an equilibrium with one another, so the hangers-on do the larger organisms no particular harm (commensalism) and may even help it (mutualism).

2. The human mouth has a plentiful and varied flora which may include many types of bacteria and even fungi (Figure 29.24).

c. Large animals have rich floras in the large intestine, over the entire skin, and in the urogenital canals.

1. Most of the organisms that live in these areas are generally harmless and some are even essential and beneficial.

2. Some bacteria occupy spaces that pathogens might otherwise occupy and their acid byproducts inhibit the growth of other organisms.

3. Every organism living in or on another is potentially an opportunist, and if the occasion arises for a microorganism to grow faster at the expense of its host, it will do so.

4. E. coli is an example of bacteria that normally lives in the human intestinal tract. If this bacteria makes its way into the human urogenital system, it can cause infection that must be treated.

d. Every pathogen has a reservoir of infection, a place where it normally resides but may not cause any disease (Figure 29.25).

1. The reservoir of infection for human disease may be in other people, in other animals, or in the soil.

2. The agent may be transmitted to the host by direct contact, through contaminated objects or food, via airborne particles, from infected large animals, and by insects or other invertebrate vectors.

3. Vectors are animals that can transmit pathogens from one host to another.

4. Rabies is a virus that can be transmitted from mammals and birds to humans via bite wounds.

e. The most notorious vectorially transmitted disease is probably plague, caused by the bacterium Yersinia pestis.

1. The Y. pestis reservoir is primarily wild rodents, although several other kinds of mammals may be infected.

2. The Y. pestis vector is primarily rat fleas.

3. The disease known as plague devastated human populations several times in recorded history.

4. The disease can now be easily controlled with modern antibiotic therapy.

f. Many diseases caused by other organisms or viruses are spread via bites from insects, ticks, or other arthropods.

1. Malaria protozoa and yellow-fever viruses are transmitted by mosquitoes, African sleeping sickness protozoa by the tsetse fly.

2. Each pathogen has a reservoir among humans, other mammals, or birds, and it uses the arthropod as a vector.

 

g. Plants, like animals, are subject to bacterial infections.

1. Plant pathogens may have a difficult time penetrating the tough, waxy walls of their hosts.

2. Nematode worms, which open holes in the plant epidermis, are notorious vectors for plant pathogens.

29.12 Some very small bacteria are intracellular pathogens.

a. Some particularly small bacteria have developed special ways of living.

1. The rickettsias are named for their discoverer, Howard T. Ricketts, who died in Mexico in 1910 from typhus fever, a rickettsial disease he was studying (Figure 29.26).

2. The rickettsias are true bacterial cells, typically about 0.3 by 1.0 µm.

3. They are intracellular parasites that can only reproduce within a host cell because they lack certain metabolic machinery.

4. Usually they infect cells of arthropods which transmit them to humans and other animals through bites.

b. Chlamydias are also very small intracellular pathogens.

1. Chlamydias are unable to metabolize glucose to make their own ATP.

2. Chlamydias are responsible for the eye infection trachoma, a frequent cause of blindness.

3. Chlamydias are also responsible for lymphogranuloma venereum, one of the lesser-known sexually transmitted diseases of humans.

29.13 Many bacteria are used in industrial processes.

a. From the beginning of civilization, humans have taken advantage of microbial activities for their food and drink.

1. Microorganisms that produce ethanol provide a substrate for others that produce vinegar.

2. Wine must be kept anaerobic to prevent the growth of bacteria such as Acetobacter, which converts the ethanol into acetic acid.

b. Before the recent invention of refrigeration, food storage presented a challenge for most people.

1. One ancient solution was salting and smoking which produces meats such as ham, bacon, and salt herring.

2. Sauerkraut and cucumbers were pickled, a preservation technique that relies on the action of lactate-producing bacteria.

3. Lactate-producing bacteria have long been used to convert milk into buttermilk, yogurt, and cheeses.

c. Linen manufacturing also depends on the growth of bacteria.

d. Bacteria with unusual metabolic capabilities have been used to manufacture some materials commercially, such as acetone.

e. Sewage treatment plants have long depended on bacteria to reduce wastes to gases (Figure 29.27).

 

D. ADDENDUM: THE VIRUSES

29.14 Viruses are not organisms.

a. Andre Lwoff, a French virologist, observed that viruses differ from organisms in several significant ways.

1. An organism is defined as being a cell or an assemblage of cells. No viruses have such a structure.

 

2. At one point in its cycle of multiplication, a virus takes the form of particles called virions, each consisting of a nucleic acid genome enclosed in a protein covering, or capsid.

3. Viruses do not grow as cells do, by enlarging and dividing, nor do they reproduce as an organism does, either sexually or asexually.

4. A virus is totally dependent on its host cell for its energy and for translating its genome into proteins.

29.15 Virions have simple, regular structures.

a. The nucleic acid and capsid of a virion form a nucleocapsid (Figure 29.28).

1. Some nucleocapsids are surrounded by a membrane, but this feature does not give the virion the properties of a cell.

b. A protective capsid can be assembled around a nucleic acid to make either a helical or a spherical structure.

1. A helical virion is made by stacking identical protein subunits that enclose the nucleic acid in an internal groove (Figure 29.29).

2. Some viruses have such a nucleocapsid enclosed in an envelope (Figure 29.30).

3. A spherical virion is made of protein subunits that form a shell around a core of nucleic acid.

4. All spherical capsids are actually icosahedrons: solids with 20 identical triangular faces. Their architecture is that of the geodesic dome invented by R. Buckminster Fuller (Figures 29.31 and Figure 29.32).

29.16 Viruses multiply in a common pattern.

a. Viruses have varied modes of infection.

1. Bacterial viruses attach to the cell surface, and their nucleic acids are transported through the cell membrane.

2. Animal viruses often enter their host cells by being phagocytized as if they were benign particles. Once inside the cell, the virus can begin to multiply.

3. Since plant cells are enclosed in protective walls, plant viruses often need vectors, such as nematode worms or insects, to break through the cell walls.

4. Viral genomes first express early genes that turn off some of the host's activities, usually stopping transcription of the host genome and translation of host messengers.

5. As replication begins, late genes of the viral genome are expressed (Figure 29.33).

b. Eventually, the infected cell falls apart and liberates many virions that have accumulated within.

1. The virions are free now to infect other cells and repeat the cycle.

2. If the new viral proteins have been added to some cell membranes, an animal's immune system can sometimes recognize the modified cells and destroy them.

29.17 Viruses have DNA or RNA genomes.

a. Viruses are classified on the basis of several factors.

1. Virions can have either a DNA or RNA genome, which may be double-stranded or single-stranded, linear or circular.

2. The nucleocapsid may be helical or spherical, and may be enveloped or naked.

3. Many bacterial viruses combine an icosahedral head with a helical tail.

b. Deoxyviruses have DNA genomes.

1. Those with naked icosahedral capsids include many minute bacteriophages; papilloma and polyoma viruses, including some that cause tumors; adenoviruses that cause respiratory diseases and tumors; and large insect viruses.

2. The enveloped icosahedral viruses are mostly herpes viruses.

3. Deoxyviruses with naked helical capsids include bacteriophages that multiply without lysing their host cells.

4. Enveloped helical deoxyviruses are pox viruses that infect humans and other animals.

c. Riboviruses are those with RNA genomes.

1. Most riboviruses with naked icosahedral capsids are plant viruses.

2. Animal viruses with naked icosahedral capsids include rhinoviruses that cause colds, polioviruses, and the virus of foot-and-mouth disease.

3. Enveloped icosahedral riboviruses are mostly arboviruses transmitted by insect vectors, including those that cause encephalitis and yellow fever.

4. The naked ribohelical viruses are almost all plant viruses.

5. The enveloped helical riboviruses cause some well-known diseases such as influenza, mumps, measles, distemper, and rabies.

d. Interactions between viruses and their hosts run the gamut from the most virulent to the most benign.

1. Some viruses coexist with their hosts in a way that doesn't necessarily kill the host and in some cases may even do the host a certain amount of good.

2. Some viruses destroy their hosts within minutes as they multiply massively.

e. Among viruses that attack humans, a wide range of symptoms and levels of severity are demonstrated.

1. Poxes are a group of viruses that usually produce nothing worse than localized skin pustules and a great deal of discomfort from itching.

2. Rhinoviruses can cause a cold that affects the respiratory system.

3. Polioviruses can destroy critical nerve cells thus paralyzing the body.

4. Hepatitis B and Epstein-Barr virus cause chronic infections that are virtually without symptoms for years.

5. Herpes simplex type 1 infects many children and persists in a latent form in ganglia in the nervous system. Years later, stress can activate the virus so it produces cold sores.

6. The extreme of virulence is probably the Marburg viruses, including the Ebola virus, which produce hemorrhagic fevers and results in death.

29.18 Viroids are unusual agents.

a. A few plant diseases have been traced to particles called viroids that are even simpler than viruses.

1. A viroid is merely a circular, single-stranded RNA molecule of 250—370 nucleotides, much smaller than the smallest viral genome.

2. After being transmitted from one plant to another mechanically or through pollen or ovules, viroids may multiply massively in the new host's cells, mostly in the nucleoli.

3. Viroids do not act as mRNAs to direct protein synthesis, and it is not known how they cause disease.

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