Evolution of Microbial Life Lectures 16 and 17 Much of the text material in the lecture notes is from our textbook, “Essential Biology with Physiology” by Neil A. Campbell, Jane B. Reece, and Eric J. Simon (2004). I don’t claim authorship. Other sources were sometimes used, and are noted. 2 Outline • • • • • • • • • • • Brief history of early life Oxygen revolution Evolution of cells Spontaneous generation and biogenesis Four-stage hypothesis of the origin of life Prokaryotic microorganisms Eukaryotic microorganisms Chemical cycling and the cycle of life Eukaryotic microorganisms Origins of multicellular life Words and terms to know 3 A brief history of early life 4 When the Earth Was Young • • • • • Our solar system formed about 4.5 billion years ago—the universe is about 13.7 billion years old. Earth’s crust began to solidify about 4 billion years ago when its interior started to cool. Prokaryotic organisms inhabited Earth as early as 3.5 billion years ago. Within the next billion years two distinct groups of prokaryotes, archaea and bacteria, diverged. These early organisms formed rapidly, leading to evolutionary development of many species. 5 Timeline Years before present Early episode 475 million Plants and fungi colonize land 570 million All major animal phyla established 1 billion First multicellular organisms 1.7 billion Oldest eukaryotic fossils 2.5 billion Accumulation of atmospheric oxygen 3.5 billion Oldest prokaryotic fossils 4.5 billion Earth formed ‘Before present’ is defined as 1950 A.D. 6 Fossilized Prokaryotes Central and southern India http://www.lisc.ernet.in The oldest prokaryotic fossils are estimated to be 3.5 billions years old. 7 Oxygen Revolution • • • • • An oxygen revolution began on our planet about 2.5 billion years ago. Photosynthetic prokaryotes that split water molecules released large amounts of oxygen gas (O2), changing the composition of the atmosphere. O2 doomed many other prokaryotic organisms because of its oxidizing, corrosive effects. A diversity of metabolic modes evolved among the surviving organisms, including cellular respiration in eukaryotic cells that uses O2 to extract energy from food. Metabolic evolution occurred during a period of almost two billion years when only prokaryotes existed. 8 Cyanobacteria http://www.unm.edu Cyanobacteria are photosynthetic form of bacteria. 9 Components of a Prokaryotic Cell http://desertfiddlekate.blogspot.com Much simpler in structure than eukaryotic cells. 10 Earliest Eukaryotic Cells Eukaryotic cells formed almost two billion years after the first prokaryotic cells—the oldest eukaryotic fossils are about 1.7 billion years old. • Eukaryotic cells evolved from prokaryotic cells that hosted even smaller prokaryotes. • The mitochondria and chloroplasts in eukaryotic cells are the evolutionary descendents of these tiny hitchhikers. • 11 Protists The earliest eukaryotic forms are protists—they are mostly microscopic and unicellular. • Protozoa and single-cell algae (such as diatoms) are a few examples of protists. • We will discuss several forms of protists found on the Earth today in this lecture. Diatoms http://www.digitalstar.net/microecologies • 12 Prokaryotic vs. Eukaryotic Cells • • • • • • Prokaryotic cells are about one-tenth the size of eukaryotic cells—their volume is about 103 smaller. Prokaryotic cells are much older in an evolutionary sense—3.5 billion vs. 1.7 billion years for eukaryotic cells. Most prokaryotes have a cell wall exterior to their plasma membranes— these walls are chemically different to the cellulose walls of plant cells. Prokaryotic cells are structurally much simpler—the internal components generally lack membranes. Prokaryotic cells also lack a membrane-enclosed nucleus—the DNA is coiled in a nucleoid region that is not partitioned from the rest of the cell. Prokaryotic cells can be compared to open warehouses while eukaryotic cells are more like modern offices partitioned into cubicles. 13 Multicellular Organisms The next great evolutionary step for eukaryotic cells was the formation of multicellular organisms that first evolved about one billion years ago. • Modern descendants include multicellular protists such as seaweeds. • Other evolutionary paths diverging from the early protists lead to fungi, plants, and animals. • 14 Cambrian Explosion The greatest diversification of animals happened during the Cambrian explosion. • The Cambrian period was the first period of the Paleozoic era beginning about 570 million years ago. • The earliest animals lived in the Pre-Cambrian seas—they diversified extensively over a span of about 10 million years in the early Cambrian period. • The major body plans of animals, known as phyla, had evolved by the end of the Cambrian period. Cambrian sea life http://www-eaps.mit.edu • 15 Colonization of Land Life was confined to aquatic environments for about the first three billion years of biological history. • Plants, in the company of fungi, began colonizing land about 475 million years ago. • Today, the roots of most plants are aided by fungi that help absorb water and minerals from the soil—without these fungi most life would not exist. • Plants transformed the terrestrial landscape and created opportunities for all life forms including plant-eating (herbivorous) animals and their predators. • 16 Colonization of Land Vertebrates (animals with a spine), in the form of amphibians, ventured out of the water. • Frogs and salamanders are descendants of air-breathing fish with fleshy fins that could support their weight on land. • Reptiles later evolved from amphibians, and birds and mammals evolved from reptiles. • 17 How did life first originate? 18 Origin of Life We may never know with certainty of how life first began on our planet. • We can start, however, with the assumption that science seeks natural causes for natural phenomena. • Artist’s depiction of single-cell organisms on the early seafloor near volcanic vents http://www.lpi.usra.edu 19 Spontaneous Generation • • • • • From the time of ancient Greek civilization, life was thought to arise from non-living matter. The process of life emerging from non-life is called spontaneous generation. The idea persisted into the mid-1800s to explain the presence of microorganisms in food. Then, Louis Pasteur demonstrated in experiments in 1862 that microbes arise from the reproduction of existing life. The life-from-life principle is known as biogenesis—since Pasteur’s work it has been extended to all forms of life. 20 Pasteur’s Experiments Could microorganisms spontaneously form and thrive in an environment not exposed to air? http://www.morning-eart.org Louis Pasteur (1822-95) http://www.darwin.museum.ru 21 Biogenesis • • • • • • Since life did not always exist how did the first organisms arise from nonliving matter? No evidence exists today for spontaneous generation, but conditions on a young Earth were very different. The primordial atmosphere contained very little O2 to oxidize and breakdown molecules. Energy sources—including volcanoes, lightning, and ultraviolet light from the sun—were more intense than today. Many biologists think it possible that physical and chemical processes in the early environment could have produced the first cells. Although there is general agreement about the outline of how life first formed, debate continues about the sequence of steps in producing the first life forms. 22 Early Energy Sources http://cas.hamptomu.edu http://www.air-and-space.com The sun’s ultraviolet spectrum http://earthobservatory.gov 23 Four-Stage Hypothesis • • • • • The first organisms may have been products of chemical changes that occurred in four stages. Stage 1: Abiotic or non-living synthesis of small organic molecules such as amino acids and nucleotides. Stage 2: Small organic molecules joined into polymers such as proteins and nucleic acids. Stage 3: Self-replicating molecules formed, eventually making genetic inheritance possible. Stage 4: The molecules formed into pre-cells, droplets with membranes that have an internal chemistry distinct from the external environment. The four-stage hypothesis leads to predictions that can be tested in the laboratory. 24 Stage 1 • • • • • Stanley Miller constructed a laboratory apparatus in 1953 that simulated atmospheric and sea conditions that may have occurred on early-Earth. Similar experiments were performed at about the same time by Harold Urey. Miller’s apparatus produced small organic (carbon backbone) molecules including amino acids essential for life—amino acids are building blocks for proteins Other scientists extended the research by varying the conditions of the artificial atmosphere and sea. Organic molecules produced in a number of experiments included all 20 amino acids, sugars, lipids, and the nucleotides found in DNA and RNA. 25 Miller-Urey Experiments Methane molecule http://classweb.smu.edu http://www.kennislink.nl Miller’s laboratory apparatus http://ucsdnews.ucsd.edu 26 Other Origins Some organic compounds found on Earth may have extraterrestrial origins. • Computer simulations have demonstrated how adenine (a molecule in DNA) may have formed in chemical reactions of cyanide in gas clouds of interstellar space. • These simulations would explain why meteorites found on earth contain organic molecules. • Meteorite on display http:www.grisda.org 27 http://www.rpi.edu Primordial Soup Add energy? 28 Stage 2 Experiments were conducted to determine if organic molecules could have formed into polymers without the assistance of enzymes found in living cells. • The synthesis of polymers has been shown to occur when organic monomers in an aqueous (water) solution are dripped onto hot sand, clay, or rock. • Heat vaporizes the water in the solution and concentrates monomers on the hot substrate. • Some of the monomers bond to form polymers, including polypeptides from amino acids. • 29 Stage 2 On early-Earth, rain may have splashed inorganic molecule solutions onto fresh lava. • These newly-formed polymers could then have been washed into the oceans. • Abiotic synthesis may have occurred near deep sea vents that emitted hot gases and superheated the water. • Deep sea vent http://fti.need.wisc.edu 30 Stage 3 The definition of life is partly based on inheritance, which is based on self-replicating molecules. • Cells store their genetic information as DNA, which is transcribed to RNA and then translated to specific proteins. • These mechanisms probably emerged through a long series of refinements to much simpler processes. • 31 First Genes? • • • • • The first genes may have been short strands of RNA that could replicate without the assistance of enzymes formed from proteins. Laboratory experiments have demonstrated that short strands of RNA can assemble spontaneously from nucleotides without the presence of enzymes. The result is a population of RNA molecules of random sequences of nucleotides monomers. On early-Earth, some of these RNA strands may have replicated with varied success resulting in molecular evolution on a very small scale. In keeping with the principles of natural selection, the RNA molecules that replicated the fastest increased their frequency in the population. 32 Stage 4 • • • • • Properties of life emerged from interaction of molecules self-organizing into higher levels of order needed to form pre-cells and then living cells. RNA strands and polypeptides may have cooperated inside a simple membrane. Physical processes needed for cellular functioning, including osmosis and voltage differences, would have been possible. Some molecular aggregates became pre-cells that contained some of the properties of life. One property is a rudimentary metabolism of absorbing molecules from the surroundings and releasing products of their reactions. 33 Chemical to Darwinian Evolution Mutations produced variations in the populations of pre-cells—the most successful pre-cells would have continued to grow, divide, and evolve. • Molecular cooperatives of pre-cells could have become more cell-like after many millions of years of refinement through the natural selection process. • The point at which pre-cells, with their rudimentary metabolism and genetic instructions, became actual living cells is not clear and may never be known. • However, life in the form of prokaryotes was in abundance 3.5 billion years ago—all other branches of life arose from these earliest cells. • 34 Prokaryotic Microorganisms 35 Prokaryotes Prokaryotic cells evolved as the only form of life on Earth for almost two billion years. • Prokaryotes have continued to adapt and flourish on an evolving Earth, and have helped to change our planet. • We will explore this diversity and adaptation in the next sections of this lecture. • 36 Where They Live Today, prokaryotes are found wherever life exists—they outnumber all eukaryotes combined. • The bacteria inhabiting a handful of fertile soil or the skin or mouth of a human can number in the billions. • Some prokaryotes thrive in habitats too hot, cold, acidic, or alkaline for eukaryotes to survive. • For example, prokaryotes were found in a goldmine almost two miles below the Earth’s surface. • 37 Bacteria in Soil http://soils.usda.gov One teaspoon of soil contains 100 million to one billion bacteria. The bacteria on an acre is equivalent to the mass of up to two cows. 38 Collective Impact Although very small organisms, prokaryotes collectively have a substantial impact on life. • The bacterial disease, bubonic plague (also known as the the black death) killed at least 25 percent of Europe’s population during the 14th century. • Tuberculosis, cholera, many sexually-transmitted diseases, and some types of food poisoning are caused by bacteria. • The Black Death http://upload.wikimedia.org 39 Bacteria on a Pin Head Electron micrograph http://faculty.ccri.edu 40 Beneficial Roles • • • • • • However, bacteria that have beneficial roles are more common than harmful bacteria. Bacteria in our intestines aid in digestion and other bacteria prevent harmful fungi from inhabiting our mouths. Bacteria that decompose dead organisms are found in the soil, and in the bottoms of rivers, lakes, and oceans. Decomposers return chemical elements to the environment in the form of inorganic compounds that can be used by plants, which in turn feed animals directly or indirectly. If decomposers were to disappear, the chemical cycles that sustain life would stop, and eukaryotic life could not survive. Prokaryotes would continue to survive as they had during the first ~ 1.7 billion years of life. 41 Prokaryotic Evolution • • • • • The five-kingdom classification is based on fundamental differences in cellular organization between prokaryotes and eukaryotes. Prokaryotes make-up the kingdom Monera separate from the eukaryotic kingdoms of Protista, Fungi, Plantae, and Animalia. The branches of prokaryotes are archaea and bacteria—the branches were established by comparing genomes. Archaea and bacteria also differ in many of their structural, biochemical, and physiological characteristics. Archaea are more related to eukaryotes than bacteria, which has led to a newer three-domain classification of Bacteria, Archaea, and Eukarya. 42 Extremophiles The majority of prokaryotes are bacteria—archaea occupy evolutionary and ecological niches. • Biologists often refer to some archaea as extremophiles, or lovers of the extreme (phile = love). • Extreme halophiles (salt lovers) thrive in the Great Salt Lake and in salt-evaporating ponds where salinity is well above that of the oceans. • Extreme thermophiles (heat lovers) live in very hot water such as deepsea vents that produce superheated water. • 43 Salt Evaporation Ponds http://fig.cox.edu Archaea thrive in high-salinity conditions, producing the yellow and red colors in the evaporation ponds. 44 Hydrothermal Sea Vents Sea vents are typically located in oceanic spreading zones http://www.is.tokayu.ac.jp The underlying hot magma superheats the seawater. http://www.mbary.org 45 Another Class of Extremophiles Some archaea live in anaerobic environments and give-off methane (CH4) as a waste product. • These methanogens living in the mud of lakes and swamps produce methane-rich marsh gas—some instances may have been resulted in UFO sightings. • http://www.freesteel.co.uk http://ugos.homestead.com 46 Other Methanogens Methanogens, most notably bacteria, inhabiting the intestine tracts of animals can produce bloating and intestinal gas. • Methanogens also help grazing animals digest the cellulose in plant walls—the animals emit large amounts of methane through belching. • http://www.classohm.com 47 Bacterial Shapes Cell shape is used as the general classification method for the many types of bacteria. • Spherical species are cocci (plural for coccus) from the Greek word for ‘berries.’ - Cocci in clusters are staphylcocci (clusters of grapes), or staph for short as in ‘staph infection.’ - Cocci in chains are streptococci (twisted grapes)—the bacterium that causes strep throat is a streptococcus. • Rod-shaped species are bacilli (plural for bacillus). • Spiral-shaped species are spirochetes—the bacterium responsible for syphilis is a spirochete. • 48 Bacterial Shapes The colors reflects the chemical stains used Bacilli Cocci Spirochetes All images from http://www.bioweb.uncc.edu 49 Cell Organization Most prokaryotes are unicellular and very small, although there are exceptions. • Some species aggregate transiently into groups of like cells, such as in streptococci. • Other prokaryotes form permanent colonies of identical, non-differentiated cells. • Some species of prokaryotes show a simple multicellular organization and divide labor between specialized types of cells. • 50 Cell Motility About half of all types of prokaryotic cells are motile, or self-propelled. • They may have one or more flagella that can propel them from unfavorable places, or favorable places such as nutrient-rich environments. • http://contents.answers.com 51 Endospores • • • • • Few bacteria can thrive in extreme environments favored by archaea. Some bacteria, however, can survive extended time periods in harsh conditions by forming resting cells, or endospores—they can remain dormant for centuries. Boiling water generally cannot kill these dormant cells. Laboratory and medical equipment are sterilized using an autoclave (similar to a pressure cooker) that kills endospores with high-pressure steam at 250o F, or 121o C. The food-canning industry uses a similar method to kill endospores of the soil bacterium Clostridium botulinum that causes botulism, a potentially fatal disease. 52 Endospore http://dwb.unl.edu 53 Anthrax Endospores http://www.sciencedaily.com Bacillus anthracis causes anthrax, an often-deadly disease in cattle, sheep, and humans. The endospores can survive harsh conditions. They can absorb water and the bacteria will resume their growth when conditions are more hospitable. 54 Reproduction Rates • • • • • • Most prokaryotes can reproduce at very high rates if conditions are favorable. The cells copy their DNA almost continuously and divide repeatedly through binary fission. If a bacteria could double every 20 minutes, in just 24 hours a single cell could give rise to a bacterial colony equivalent in mass to 15,000 humans. Bacteria cannot sustain exponential growth due to limited resources including food and space, and the waste products from the bacteria that pollute the colony’s environment. The rapid and repeated doubling helps to explain why a few bacteria cells can make us very ill. Refrigeration (cold) retards food spoilage since most microorganisms reproduce very slowly at low temperatures. 55 Repeated Geometric Doubling 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 1 2 4 8 16 32 64 128 256 512 1,024 2,048 4,096 8,192 16,384 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 32,768 65,536 131,072 262,144 524,288 1,048,576 2,097,152 4,194,304 8,388,608 16,777,216 33,554,432 67,108,864 134,217,728 268,435,456 536,870,912 Would you rather have a million dollars or a dollar that doubles every day for 30 days? 56 Nutritional Diversity • • • • • The prokaryotic domain has a variety of biological mechanisms for obtaining nutrition. Nutrition refers to how an organism obtains energy and carbon for synthesizing organic compounds. Phototrophs use light energy; chemotrophs obtain their energy from their environment; autotrophs need only carbon dioxide for a carbon source. Heterotrophs require at least one organic nutrient (such as glucose) as a carbon source. All prokaryotes can be grouped according to four modes of nutrition as shown on the next slide. 57 Modes of Nutrition Nutritional Type Energy Source Carbon Source Examples Photoautotroph (photosy nthesizer) Sunlight CO 2 Cyanobacter ia (similar mechanism to photosynthesis in plants) Chemo autotroph Inorga nic chemicals CO 2 Prokaryotes living near deep-water heat vents Photoheterotroph Sunlight Organic compounds Very limited number of prokaryotes Organic compounds Widely used among prokaryotes (a nd animals and fungi too) Chemo heterotroph Organic compounds 58 Bacteria that Cause Disease • • • • • We are constantly exposed to bacteria, and some species be harmful. Bacteria and other microorganisms that cause diseases are known as pathogens. The body’s defenses, including the immune system, usually inhibit the growth of pathogens. Sometimes the balance shifts in the favor of the pathogen, and illness results. Some of the bacteria that are normally residents in our bodies can make us ill when our defenses are weakened by a viral infection or poor nutrition. 59 H. Influenzae Bacterium The Haemophilus influenzae bacterium resides on the epithelial cells of the interior of the nose, and can cause pneumonia. • Malnourished children are particularly susceptible to this bacterium due to their lowered resistance. • This bacterium, although similar in name, is not the same as the flu virus. • http://www.brown.edu 60 Exotoxins • • • • • Pathogenic bacteria cause disease by producing poisons—they can be either exotoxins or endotoxins. Exotoxins are proteins secreted by bacterial cells—a single gram of botulism could kill one million people if it were widely dispersed. The exotoxin producer, Staphylococcus aureus, is a usually harmless resident of the surface of human skin. Serious disease can result if this bacteria (known as staph) enters the body through a wound or contaminated food. S. aureus can produce a variety of effects including sloughing of skin layers, vomiting, severe diarrhea, and toxic shock syndrome, which can be deadly. 61 Endotoxins Endotoxins are chemical components of cell walls of some bacteria. • All endotoxins produce the same general set of symptoms—fever, aches, and shock (a dangerous drop in blood pressure). • The severity of these conditions varies with the host organism and the type of bacteria. • Different species of the bacteria Salmonella produce endotoxins that cause food poisoning and typhoid fever. • 62 Prevention and Treatment • • • • • Bacterial infections have progressively decreased in many countries during the past century following the discovery that microorganisms cause disease. Bacterial infections can be addressed through sanitation, antibiotics, and education. Sanitation methods are the most effective ways to prevent bacterial disease. Installation and operation of water treatment and sewage systems is an effective public health approach. Antibiotics are used to treat most bacterial diseases, but their widespread use has resulted in the evolution of drug-resistant pathogens, such as some forms of tuberculosis. 63 Lyme Disease • • • • • • Lyme disease is caused by a spirochete bacterium carried by ticks found on deer and field mice. It is the most widespread pest-carried disease in the United States, especially in the mid-Atlantic and northeastern states (some cases have been reported on the West Coast). Early symptoms of Lyme disease is a red rash shaped like a bull’s eye encircling a tick bite. Antibiotics can be used in effectively treating the disease if they are given within a month of exposure. If left untreated, Lyme disease can cause debilitating arthritis, heart disease, and disorders of the nervous system. The best defense is public education about using insect repellant, wearing long sleeve shirts and long pants, and seeking treatment if a characteristic rash develops. 64 http://aapredbook.aappublications.org Lyme Disease http://www.lib.uiowa.edu http://www.nlm.nih.gov 65 Chemical Cycling • • • • • All life depends on the recycling of chemical elements between the biological and physical components of ecosystems. Cyanobacteria restore oxygen to the atmosphere and some convert nitrogen gas in the atmosphere to nitrogen compounds that plants can then absorb from soil and water. Bacteria living in the roots of bean plants and other legumes contribute large amounts of nitrogen to the soil. All nitrogen that plants use in synthesizing proteins and nucleic acids comes from prokaryotic metabolism. As consumers, animals must obtain their nitrogen compounds from plants. 66 The Cycle of Life Some prokaryotes break down organic waste and dead organisms. • Through this process of decomposition, prokaryotes return elements in inorganic forms that can be used by other organisms. • Carbon, nitrogen, and other essential elements would be locked in waste products and dead organisms if not for decomposers such as prokaryotes. • Life would eventually cease to exist if these essential elements could not be used. • 67 Bioremediation Biologically-diverse prokaryotes are being used in helping to cleanup the environment. • Use of microorganisms to remove pollutants from water, air, and soil is known as bioremediation. • http://www.alabastercorp.com 68 Abandoned Mine Site Rocky Mountains, Colorado http://toxics.usgs.gov Many long-abandoned mines continue to be toxic waste hazards. 69 Bioremediation • • • • • Anaerobic prokaryotes are added to solid matter (sludge) produced from the raw effluent at sewage treatment plants. The microorganisms decompose the organic matter in the sludge for use as landfill or as fertilizer after chemical sterilization. Bacteria that naturally occur on beaches can decompose petroleum and help clean-up oil spills. Genetically-engineered bacteria in development to accelerate the oil-consuming process. Other bacteria are being genetically-engineered to consume and deactivate poisonous heavy metals found at old, often abandoned mining sites. 70 Eukaryotic Microorganisms 71 Pond Water http://www.georgeloper.org Walden Pond, Concord, Massachusetts http://www.visitusa.com 72 Protists Pond water observed under a light microscope reveals many diverse organisms known as protists. • Protists are eukaryotes—even the simplest are more complex than prokaryotic cells. • The first eukaryotes composed of single-cell organisms evolved from prokaryotic cells—multicellular eukaryotes were to follow many millions of years later. • 73 Origin of Eukaryotic Cells • • • • • • Differences between prokaryotic and eukaryotic cells far outnumber the differences between animal and plant cells. Fossil records suggest the first eukaryotes evolved from prokaryotes about 1.7 billion years ago. A major question, yet to be fully answered, is how eukaryotes first emerged. One theory is eukaryotic cells emerged through the combination of two processes. In the first process the endomembrane system formed from inward folding of the plasma membrane in prokaryotic cells. A second process, endosymbiosis, produced early mitochondria and chloroplasts. The endomembrane system includes all membrane-enclosed cells in the cytoplasm except for the mitochondria and chloroplasts. 74 Endosymbiosis Symbiosis is a close association between organisms of two or more species where there is mutual benefit. • Endosymbiosis refers to a species living within a host species. • Mitochondria and chloroplasts probably evolved from smaller prokaryotes that established residence within larger host prokaryotic cells. • 75 Origin of Mitochondria and Chloroplasts The ancestors of mitochondria may have been aerobic bacteria that used oxygen to generate energy in the form of ATP and NADPH. • Some of the bacteria may have escaped digestion by the larger cell and continued to perform respiration in support of its host as oxygen accumulated in the atmosphere. • Photosynthetic bacteria may have also lived in host cells, and later evolved into chloroplasts. • Mitochondria may have developed first since all eukaryotic cells have mitochondria, but not all have chloroplasts. • 76 Similarity with Prokaryotic Cells • • • • • Mitochondria and chloroplasts are similar to prokaryotic cells in key ways. Both contain DNA, RNA, and ribosomes that resemble prokaryotic versions. These genetic components enable mitochondria and chloroplasts to show some autonomy. For example, they transcribe and translate their DNA into polypeptides to synthesize their own enzymes. Mitochondria and chloroplasts also replicate their DNA by a process resembling binary fission in prokaryotes. 77 Wide Diversity of Protists • • • • • • Although all protists are eukaryotes they are so diverse they share few other general characteristics. Protists vary in structure and function more than any other group of organisms. Most protists are unicellular, but some are colonial or multicellular. Protists are the most complex of all cells since unicellular protists must be as complete as any plant or animal to perform the functions necessary for life. The four major categories of protists are: protozoans, slime mold, unicellular algae, and seaweeds. The categories are organized more by lifestyle than by evolutionary relationships. 78 Star Wars Cantina Scene http://www.starwarsholidayspecial.com The diverse world of protists reminds me of this movie scene. 79 Protozoans • • • • • Protists that obtain nourishment by ingesting food are protozoans, or ‘first animals.’ They thrive in all types of aquatic environments including lakes, wet soil, and the fluid environments inside other organisms. Most species eat bacteria—some can absorb nutrients dissolved in water. Protozoans living as parasites in animals, and cause some of the world’s most harmful diseases. Protozoans include flagellates, amoebas, forams, apicomplexans, and ciliates. 80 Flagellates • • • • • • This group of protozoans locomote by the means of one or more flagella. Most flagellates are free-living and usually not dangerous, but some are parasitic. Giardia is a parasite that infects the human intestine and can cause abdominal cramping and severe diarrhea. This parasite can be transmitted to humans through drinking water contaminated with feces from animals. Other flagellates that are dangerous are the trypanosomes including a species that causes sleeping sickness. Sleeping sickness is a debilitating sickness found in parts of Africa; it is transmitted by the tsetse fly. 81 Prokaryotic Flagella Connecting mechanism http://images.encarta.msn.com Multiple flagella http://www.sinisia.web1000.com 82 Mountain Stream http://www.markbsplace.net Cool, clear water… http://ehrenfest.anu.edu Please be sure to filter it before drinking! 83 Amoeba Although amoeba have considerable flexibility and can move rapidly, they no permanent locomotor organelles. • Amoeba move and feed by pseudopodia—extensions of the cell resulting from rapid reconfiguration of protein microtubules in the cytoplasm. • Amoeba can assume many shapes as they move among the rocks, sticks, mud, and other obstacles in ponds, lakes, rivers, and oceans. • 84 Amoeba Feeding Reproduction (binary fission) http://www.scientificillustrator.com Images from http://www.biology-resources.com 85 Amoebic Diseases Many species of amoeba are not dangerous to humans—however, one species can enter the brain through the nasal cavity which can lead to death in a few weeks. • In the past few years some fatalities have been reported for swimmers—especially children—in the warm lakes of the southeastern United States. • Amoebic dysentery results from an amoebic parasite, which sometimes occurs in countries with poor sanitation. • 86 Forams Most forams are marine organisms—that is, they live in the world’s oceans. • The cells secrete a porous shell made of organic material hardened with calcium carbonate. • Thin strands of cytoplasm (pseudopodia) extend through the pores of the shell for swimming, feeding, and shell formation. • The shells of fossilized forams form the limestone rocks found in some land formations. • Limestone distribution (shown in yellow) http://www.reec.nsw.edu 87 Limestone Fossils embedded in limestone Both images from http://upload.wikimedia.org Limestone cliffs 88 Apicomplexans • • • • • All apicomplexans are parasitic, and some can cause human serious diseases. They are named for an apparatus at their apex (apical complex) that penetrates host cells and tissues. The apiocomplexan, Plasmodium, causes malaria that infects more than 200 million people in the tropics each year. The apical complex is used to enter and feed upon red blood cells of the human host, eventually destroying them. Malaria is spread by a few species of mosquitoes—and it is one of the most debilitating and widespread human diseases throughout history. 89 http://www.oucom.ohiou.edu Plasmodium (Malaria) Cycle 90 Malaria Map http://www.ch.ic.ac.uk High-risk areas are shown in red. 91 Ciliates Ciliates use locomotor structures made of proteins, known as cilia, to move and feed. • Most are free-living and not parasitic, and therefore pose very little danger. • The best known example of a ciliate is Paramecium, found in pond water. • http://www.silkentent.com 92 Paramecium http://en.wikivisual.com http://www.cartage.org.lb 93 More Ciliates http://www.nsf.gov 94 Slime Molds Slime molds resemble fungi in appearance and ‘lifestyle’ although they are not closely related—the similarities are due to convergent evolution. • Slime molds have filament-like bodies suited for their role of decomposers of organic material. • The two main categories of slime molds are plasmodial and cellular. • 95 Plasmodial Slime Mold Plasmodial slime molds form an amoeboid mass, known as plasmodium, during the feeding stage in their lifecycle. • Plasmodial slime molds are found among leaves and other decaying material on forest floors. • The plasmodium, a single cell with many nuclei, can measure several centimeters across. http://www.ppdl.purdue.edu • 96 Cellular Slime Mold Cellular slime molds consist of amoeboid-like cells functioning individually to feed on decaying organic matter. • When the food source is depleted the individual cells aggregate and form a slug-like colony that moves and functions as a single unit. • Cellular slime molds raise the question about what it means to be an individual organism. • 97 Unicellular Algae • • • • • Photosynthetic protists are called algae—their chloroplasts support food chains in freshwater and marine ecosystems. Many unicellular algae are components of plankton (from the Greek word for ‘wanderer’). Plankton are communities of mostly microscopic organisms that drift or swim near the surfaces of ponds, lakes, and oceans. More specifically, these types of algae are known as phytoplankton due to their photosynthetic capabilities. Three groups of unicellular algae are dinoflagellates, diatoms, and green algae. 98 Dinoflagellates • • • • • Dinoflagellates are photosynthetic protists—they are commonly-found types of phytoplankton. Each species has a characteristic shape reinforced by external plates made of cellulose. The beating of the two flagella in perpendicular grooves produces the spinning motion for which they are named (‘dino’ means ‘whirling’ in Greek). Population explosions of dinoflagellates cause a phenomenon in warm coastal waters known as red tide. Toxins produced by these dinoflagellates can result in massive fish kills, and can be dangerous to humans too—public health alerts are issued when red tides are expected. 99 Dinoflagellates http://cache.eb.com http://bioweb.uncc.edu 100 Diatoms Diatoms have clear cell walls containing silca, the mineral used to make glass—these protists come in many shapes and sizes. • The glass-like cell walls consist of two halves that fit together as with the bottom and lid of a shoe box. • Diatoms store food reserves in oil that provides buoyancy, enabling these phytoplankton to float near the sunlit surface. • Very large accumulations of fossilized diatoms make-up thick sediments known as diatomaceous earth, which is mined for use as filtering material and an abrasive. • 101 Multitude of Diatomic Shapes http://www.botany.hawaii.edu http://www.micographia.com 102 Centric Diatoms http://www.nh.ac.uk 103 Diatomaceous Earth http://epod.usra.edu.de http://www.ucmp.berkeley.edu http://www.invisiblegardener.com 104 Green Algae • • • • • Green algae, named for their grass-green chloroplasts, flourish in many freshwater lakes and ponds. Some species are flagellated to allow for movement in their through the watery medium. Green algae also include colonial forms, such as Volvox—each colony is a ball of flagellated cells. The balls-within-balls are daughter colonies that will be released when the parent colonies rupture. Of all the photosynthetic protists, green algae are most closely related to terrestrial plants. 105 Green Algae http://www.virtualviz.com Freshwater pond. 106 Volvox http://www.rbgsyd.nsw.gov.au Close-up of a single colonial form on the right. 107 Seaweeds Seaweeds are large, multicelullar marine algae that grow on rock shores and offshore beyond the zone of pounding surf. • The cell walls contain a rubbery substance that cushions seaweed from the agitation of waves. • Although seaweeds can be as large and complex as terrestrial plants, the similarities are a consequence of convergent evolution. • The closest relatives of seaweeds are unicellular algae, which is why they are classified as protists. • 108 Human Diets • • • • • Coastal peoples, particularly in Asia, harvest seaweed as food—in Japan and Korea some species are added to soups and others to wrap sushi. Marine algae are a source of iodine, to help prevent goiter, and other minerals too. Most of the organic material in seaweeds are polysaccharides that cannot be digested by humans—seaweeds are eaten mostly for their rich tastes and textures. The gel-forming substances in the cell walls are used as thickeners in some processed foods such as puddings, ice cream, and salad dressings. Agar, a seaweed extract, provides the gelatinous base for the culture of bacteria in Petri dishes. 109 Green and Red Algae Seaweeds are classified into three groups (green, red, and brown) in part based on the types of pigments present in their chloroplasts. • Green algae often inhabit the intertidal zone between high and low tides— they are related to unicellular and colonial species of algae. • Red algae are most abundant in the warm waters of the tropics, and live in relatively deep water. • The chloroplasts in red algae have pigments that absorb blue and green light that penetrates best through water. • 110 Types of Algae Red algae Brown algae Green Algae All images from http://taggart.glg.msu.edu 111 Brown Algae • • • • • • Brown algae include the largest seaweeds, known as kelp—kelp beds are found along the coast from the Gulf of Alaska to Baja California, and near the islands in the Southern California Bight. Some species such as giant kelp can grow linearly over a foot per day and almost 200 feet in a single year. Kelp provides rich habitat for many marine species—in some locations the estimate is over 700 species. Kelp beds also calm the ocean surface and reduce the pounding of waves that can erode a shoreline. Piles of kelp can often be found on Southern California beaches especially after a storm--the small bladders are gas-filled floats that keep the photosynthetic blades near the ocean’s surface. Kelp, harvested by boats that cut and collect their tops, can be a renewable resource if managed properly. 112 Kelp http://www.onr.navy.mil Giant kelp, Macrocystis pyrifera Monterey Bay Aquarium http://upload.wikimedia.org 113 Kelp Harvesting Republic of Ireland http://www.maritimes.ie Icelandic kelp harvester Northern California http://www.thorverk.is http://montereybay.noaa.gov 114 Origins of Multicellular Life 115 Cellular Complexity • • • • • An orchestra can play a greater variety of musical compositions than a violin soloist can. The general concept is that the greater complexity makes more variations possible. The origin of the eukaryotic cell led to an evolutionary ‘radiation’ of new life forms. Unicellular protists, which are eukaryotic cells, are more diverse in form than the simpler prokaryotic cells. The evolution of multicelullar complexes of eukaryotic cells broke through an even higher threshold in structural organization. 116 Multicellular vs. Unicellular Organisms Multicellular and unicellular organisms are fundamentally different from one other. • All of life’s activities occur within the single cell of a unicellular organism. • A multicelullar organism, in comparison, has specialized cells that perform different functions and are dependent on each other. • For example some cells procure food, others transport materials, and yet others enable movement. • 117 Evolutionary Links • • • • • The evolutionary links between unicellular and multicellular organisms were may have been colonial forms in which unicellular forms joined together to form federations of independent cells. The gradual transition from colonies to multicellular organisms involved cells becoming increasingly more interdependent as divisions of labor evolved. Individual cells in multicellular organisms are specialized to perform many functions including reproduction, feeding, waste disposal, gas exchange, and protec-tion, among others. Multicellularity may have evolved many times among ancestral protists, leading to new waves of biological diversification and species that are found today. Seaweeds, plants, fungi, and animals are all descendents of protists on a much younger Earth. 118 Words and Terms to Know • • • • • • • • • • • • • • • • Algae Amoeba Apicomplexans Archaea Bacteria Bacterial shapes Biogenesis Bioremediation Chemical cycling Ciliates Convergent evolution Cyanobacteria Decomposers Diatoms Dinoflagellates Endospores•••••••••••••• Endotoxins and exotoxins • Exotoxins • Extremophiles • Eukaryotic cells • Flagella • Forams • Four-stage hypothesis • Lyme disease • Metabolic evolution • Microorganisms • Plankton • Prokaryotic cells• • Protists • Protozoans • Slime molds • Spontaneous generation • 119 Miller’s Laboratory Apparatus http://upload.wikimedia.org 120