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MAJOR EPISODES IN THE HISTORY OF LIFE• Earth was formed about 4.6 billion years ago. • Prokaryotes
– evolved by about 3.5 billion years ago, – began oxygen production about 2.7 billion years
ago, – lived alone for more than a billion years, and – continue in great abundance today.
– Eukaryotes • Single-celled eukaryotes first evolved about 2.1 billion years ago. • Multicellular eukaryotes first evolved at least 1.2 billion years ago.
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Figure 15.1a
Precambrian
Ancestor to all present-day life
Origin of Earth
Earth’s crust solidifies
Oldest prokaryotic fossils
Atmospheric oxygen begins to appear
Millions of years ago4,500 4,000 3,500 3,000 2,500
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Figure 15.1b
Millions of years ago2,000 1,500 1,000
Precambrian
Oldest eukaryotic fossils
Origin of multicellular organisms
Oldest animal fossils
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Figure 15.1c
Millions of years ago
1,000 500 0
Precambrian Paleozoic CenozoicMeso- zoic
Bacteria
Archaea
PlantsFungi
AnimalsCambrian explosion
Oldest animal fossils
Plants colonize land
Extinction of dinosaurs
First humans
Protists
Prokaryotes
Eukaryotes
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• All the major phyla of animals evolved by the end of the Cambrian explosion, which – began about 540 million years ago and – lasted about 10 million years.
• Plants and fungi – first colonized land about 500 million years ago and – were followed by amphibians that evolved from fish.
MAJOR EPISODES IN THE HISTORY OF LIFE
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• What if we use a clock analogy to tick down all of the major events in the history of life on Earth?
MAJOR EPISODES IN THE HISTORY OF LIFE
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Figure 15.2
Humans
Origin of solar system and Earth
1 4
2 3
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Figure 15.2a
Origin of solar system and Earth
1 4
0
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Pr entes
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Figure 15.2b
2 3
Sing
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THE ORIGIN OF LIFE
• We may never know for sure how life on Earth began.
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Resolving the Biogenesis Paradox
• All life today arises by the reproduction of preexisting life, or biogenesis. • If this is true, how could the first organisms arise? • From the time of the ancient Greeks until well into the 1800s, it was commonly
believed that life regularly arises from nonliving matter, an idea called spontaneous generation.
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• Today, most biologists think it is possible that life on early Earth evolved from simple cells produced by
– chemical and – physical processes.
Resolving the Biogenesis Paradox
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Figure 15.3
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A Four-Stage Hypothesis for the Origin of Life
• According to one hypothesis, the first organisms were products of chemical evolution in four stages.
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Stage 1: Abiotic Synthesis of Organic Monomers
• The first stage in the origin of life was the first to be extensively studied in the laboratory.
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The Process of Science: Can Biological Monomers Form Spontaneously?• Observation: Modern biological macromolecules are all composed of elements
that were present in abundance on early Earth. • Question: Could biological molecules arise spontaneously under conditions like
those on early Earth?
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• Hypothesis: A closed system designed to simulate early Earth conditions could
produce biologically important organic molecules from inorganic ingredients. • Prediction: Organic molecules would form and accumulate.
The Process of Science: Can Biological Monomers Form Spontaneously?
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• Experiment: An apparatus was built to mimic the early Earth atmosphere and included
– hydrogen gas (H2), methane (CH4), ammonia (NH3), and water vapor (H2O),
– sparks that were discharged into the chamber to mimic the prevalent lightning of early Earth, and
– a condenser that cooled the atmosphere, causing water and dissolved compounds to “rain” into the miniature “sea.”
The Process of Science: Can Biological Monomers Form Spontaneously?
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Figure 15.4
Miller and Urey’s experiment
“Sea”
H2O
Sample for chemical analysis
Cooled water containing organic molecules
Cold water
Condenser
Electrode
“Atmosphere”Water vapor
CH4
NH3 H2
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• Results: After the apparatus had run for a week, an abundance of organic molecules essential for life had collected in the “sea,” including amino acids, the monomers of proteins.
• These laboratory experiments – have been repeated and extended by other
scientists and – support the idea that organic molecules could have
arisen abiotically on early Earth.
The Process of Science: Can Biological Monomers Form Spontaneously?
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Stage 2: Abiotic Synthesis of Polymers
• Researchers have brought about the polymerization of monomers to form polymers, such as proteins and nucleic acids, by dripping solutions of organic monomers onto – hot sand, – clay, or – rock.
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Stage 3: Formation of Pre-Cells
• A key step in the origin of life was the isolation of a collection of abiotically created molecules within a membrane.
• Laboratory experiments demonstrate that pre-cells could have formed spontaneously from abiotically produced organic compounds.
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• Such pre-cells produced in the laboratory display some lifelike properties. They – have a selectively permeable surface, – can grow by absorbing molecules from their
surroundings, and – swell or shrink when placed in solutions of different
salt concentrations.
Stage 3: Formation of Pre-Cells
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Stage 4: Origin of Self-Replicating Molecules
• Life is defined partly by the process of inheritance, which is based on self-replicating molecules.
• One hypothesis is that the first genes were short strands of RNA that replicated themselves
– without the assistance of proteins, – perhaps using RNAs that can act as enzymes,
called ribozymes.
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Figure 15.5-1
RNA monomers
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Figure 15.5-2
RNA monomers
Formation of short RNA polymers: simple “genes”
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Figure 15.5-3
RNA monomers
Formation of short RNA polymers: simple “genes”
Assembly of a complementary RNA chain
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Figure 15.5-4
Original “gene”
RNA monomers
Formation of short RNA polymers: simple “genes”
Assembly of a complementary RNA chain
The complementary chain serves as a template of the original “gene.”
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From Chemical Evolution to Darwinian Evolution
• Over millions of years – natural selection favored the most efficient pre-cells
and – the first prokaryotic cells evolved.
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PROKARYOTES
• Prokaryotes lived and evolved all alone on Earth for about 2 billion years.
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Vox Bacteria Video
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They’re Everywhere!
• Prokaryotes – are found wherever there is life, – have a collective biomass that is at least ten times
that of all eukaryotes, – thrive in habitats too cold, too hot, too salty, too
acidic, or too alkaline for any eukaryote, – cause about half of all human diseases, and – are more commonly benign or beneficial.
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Figure 15.6
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• Compared to eukaryotes, prokaryotes are – much more abundant and – typically much smaller.
They’re Everywhere!
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Figure 15.7
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• Prokaryotes living in soil and at the bottom of lakes, rivers, and oceans help to decompose dead organisms and other organic waste material, returning vital chemical elements to the environment.
They’re Everywhere!
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The Structure and Function of Prokaryotes
• Prokaryotic cells – lack a membrane-enclosed nucleus, – lack other membrane-enclosed organelles, – typically have cell walls exterior to their plasma
membranes, but – display an enormous range of diversity.
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Figure 4.4
Plasma membrane
Cell wall
Capsule
Prokaryotic flagellum
Ribosomes
Nucleoid
Pili
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Figure 4.5
CytoskeletonRibosomes Centriole
Lysosome
Nucleus
Plasma membrane
Mitochondrion
Rough endoplasmic reticulum (ER)
Golgi apparatus
Smooth endoplasmic reticulum (ER)
Idealized animal cell
Idealized plant cell
CytoskeletonMitochondrion
Nucleus
Rough endoplasmic reticulum (ER)
Ribosomes
Smooth endoplasmic
reticulum (ER)
Golgi apparatus
Plasma membrane
Channels between cells
Not in most plant cells
Central vacuoleCell wallChloroplast
Not in animal cells
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• The three most common shapes of prokaryotes are 1. spherical (cocci), 2. rod-shaped (bacilli), and 3. spiral or curved.
Prokaryotic Forms
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Figure 15.8
SHAPES OF PROKARYOTIC CELLSSpherical (cocci) Rod-shaped (bacilli) Spiral
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Figure 15.8a
Spherical (cocci)
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Figure 15.8b
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Rod-shaped (bacilli)
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Figure 15.8c
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Spiral
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• All prokaryotes are unicellular. • Some species
– exist as groups of two or more cells, – exhibit a simple division of labor among specialized
cell types, or – are very large, dwarfing most eukaryotic cells.
Prokaryotic Forms
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Figure 15.9
(a) Actinomycete
(b) Cyanobacteria (c) Giant bacteriumC
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• About half of all prokaryotes are mobile, and many of these travel using one or more flagella.
Prokaryotic Forms
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• In many natural environments, prokaryotes attach to surfaces in a highly organized colony called a biofilm, which – may consist of one or several species of
prokaryotes, – may include protists and fungi, – can show a division of labor and defense against
invaders, and can form on almost any type of surface, including
– rocks, – metal, – plastic, and – organic material including teeth.
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Figure 15.10
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• Most prokaryotes can reproduce – by dividing in half by binary fission and – at very high rates if conditions are favorable.
• Some prokaryotes form endospores, which are – thick-coated, protective cells – produced when the prokaryote is exposed to
unfavorable conditions.
Prokaryotic Reproduction
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Figure 15.11
Endospore
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Prokaryotic Nutrition
• Biologists use the phrase “mode of nutrition” to describe how organisms obtain energy and carbon. – Energy
– Phototrophs obtain energy from light. – Chemotrophs obtain energy from environmental
chemicals.
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Prokaryotic Nutrition
– Carbon – Autotrophs obtain carbon from carbon dioxide
(CO2). – Heterotrophs obtain carbon from at least one
organic nutrient—the sugar glucose, for instance.
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• We can group all organisms according to the four major modes of nutrition if we combine the – energy source (phototroph versus chemotroph) and – carbon source (autotroph versus heterotroph).
Prokaryotic Nutrition
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• Dominant among multicellular organisms are – photoautotrophs and – chemoheterotrophs.
• The other two modes are used only by certain prokaryotes.
Prokaryotic Nutrition
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Figure 15.12
MODES OF NUTRITION
Light ChemicalChemoautotrophsPhotoautotrophs
Photoheterotrophs Chemoheterotrophs
Energy source
Elodea, an aquatic plant
Rhodopseudomonas Kingfisher with prey
Bacteria from a hot spring
Org
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Figure 15.12a
A photoautotroph: Elodea, an aquatic plant
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Figure 15.12b
A chemoautotroph: Bacteria from a hot spring
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Figure 15.12c
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A photoheterotroph: Rhodopseudomonas
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Figure 15.12d
A chemoheterotroph: Kingfisher with prey
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The Two Main Branches of Prokaryotic Evolution: Bacteria and Archaea• By comparing diverse prokaryotes at the molecular level, biologists have
identified two major branches of prokaryotic evolution: 1. bacteria and 2. archaea (more closely related to eukaryotes).
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The Two Main Branches of Prokaryotic Evolution: Bacteria and Archaea• Thus, life is organized into three domains:
1. Bacteria, 2. Archaea, and 3. Eukarya.
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• Some archaea are “extremophiles.” – Halophiles thrive in salty environments. – Thermophiles inhabit very hot water. – Methanogens
– inhabit the bottoms of lakes and swamps and – aid digestion in cattle and deer.
The Two Main Branches of Prokaryotic Evolution: Bacteria and Archaea
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Figure 15.13
(a) Salt-loving archaea (b) Heat-loving archaea
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Bacteria and Disease Bacteria That Cause Disease• Bacteria and other organisms that cause disease are called pathogens. • Most pathogenic bacteria produce poisons.
– Exotoxins are proteins bacterial cells secrete into their environment.
– Endotoxins are – not cell secretions but instead – chemical components of the outer membrane of
certain bacteria.
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Figure 15.14
Haemophilus influenzae
Cells of nasal lining
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• The best defenses against bacterial disease are – sanitation, – antibiotics, and – education.
Bacteria That Cause Disease
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• Lyme disease is – caused by bacteria carried by ticks and – treated with antibiotics, if detected early.
Bacteria That Cause Disease
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Figure 15.15
“Bull’s-eye” rash
Tick that carries the Lyme disease bacterium
Spirochete that causes Lyme disease
SEM
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The Ecological Impact of Prokaryotes
• Pathogenic bacteria are in the minority among prokaryotes. • Far more common are species that are essential to our well-being, either
directly or indirectly.
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• Prokaryotes play essential roles in – chemical cycles in the environment and – the breakdown of organic wastes and dead
organisms.
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Prokaryotes and Bioremediation
• Bioremediation is the use of organisms to remove pollutants from – water, – air, and – soil.
• A familiar example is the use of prokaryotic decomposers in sewage treatment.
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Figure 15.17
Liquid wastes Outflow
Rotating arm spraying liquid wastes
Rock bed coated with aerobic prokaryotes and fungi
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• Certain bacteria – can decompose petroleum and – are useful in cleaning up oil spills.
Prokaryotes and Bioremediation
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Figure 15.18
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PROTISTS
• Protists are – eukaryotes that are not fungi, animals, or plants, – mostly unicellular, and – ancestral to all other eukaryotes.
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The Origin of Eukaryotic Cells
• Eukaryotic cells evolved by – the infolding of the plasma membrane of a
prokaryotic cell to form the endomembrane system and
– a process known as endosymbiosis.
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The Origin of Eukaryotic Cells
• Symbiosis is a more general association between organisms of two or more species.
• Endosymbiosis – refers to one species living inside another host
species and – is the process by which eukaryotes gained
mitochondria and chloroplasts.
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Figure 15.19
(a) Origin of the endomembrane system (b) Origin of mitochondria and chloroplasts
Plasma membraneDNA
Cytoplasm
Endoplasmic reticulum
Nucleus
Nuclear envelope
Ancestral prokaryote
Membrane infolding
Cell with nucleus and endomembrane system
Photosynthetic eukaryotic cell
Photosynthetic prokaryote
Aerobic heterotrophic prokaryote
Endosymbiosis
Mitochondrion
Chloroplast
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Figure 15.19a
(a) Origin of the endomembrane system
Plasma membraneDNA
Cytoplasm
Endoplasmic reticulum
Nucleus
Nuclear envelope
Ancestral prokaryote
Membrane infolding
Cell with nucleus and endomembrane system
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Figure 15.19b
(b) Origin of mitochondria and chloroplasts
Photosynthetic eukaryotic cell
Photosynthetic prokaryote
Aerobic heterotrophic prokaryote
Endosymbiosis
Mitochondrion
Chloroplast
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The Diversity of Protists
• The group called protists – consists of multiple clades but – remains a convenient term to refer to eukaryotes
that are not plants, animals, or fungi.
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• Protists obtain their nutrition in a variety of ways. – Algae are autotrophs, producing their food by
photosynthesis.
The Diversity of Protists
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– Other protists are heterotrophs. – Some protists eat bacteria or other protists. – Other protists are fungus-like and obtain organic
molecules by absorption. – Parasites derive their nutrition from a living host,
which is harmed by the interaction. Parasitic trypanosomes infect blood and cause sleeping sickness.
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– Other protists are heterotrophs. – Some protists eat bacteria or other protists. – Other protists are fungus-like and obtain organic
molecules by absorption. – Parasites derive their nutrition from a living host,
which is harmed by the interaction. Parasitic trypanosomes infect blood and cause sleeping sickness.
The Diversity of Protists
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Figure 15.20
(a) An autotroph: Caulerpa, a multicellular alga
(b) A heterotroph: parasitic trypanosome
(c) A mixotroph: Euglena
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• Protist habitats are diverse and include – oceans, lakes, and ponds, – damp soil and leaf litter, and – the bodies of host organisms with which they share
mutually beneficial relationships, such as – unicellular algae and reef-building coral animals,
and – cellulose-digesting protists and termites.
The Diversity of Protists
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Figure 15.21
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Protozoans
• Protists that live primarily by ingesting food are called protozoans.
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Figure 15.22
A foram
A flagellate: Giardia
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Another flagellate: Trichomonas An amoeba
Red blood cell
Apical complex
TEM
An apicomplexan A ciliate
Cilia
Cell “mouth”
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• Protozoans with flagella are called flagellates and – are typically free-living, but – some are nasty parasites.
Protozoans
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Figure 15.22a
A flagellate: Giardia
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• Amoebas are characterized by – great flexibility in their body shape and – the absence of permanent organelles for
locomotion. • Most species move and feed by means of pseudopodia (singular,
pseudopodium), temporary extensions of the cell.
Protozoans
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Figure 15.22c
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An amoeba
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• Other protozoans with pseudopodia include the forams, which have shells.
Protozoans
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• Apicomplexans are – named for a structure at their apex (tip) that is
specialized for penetrating host cells and tissues, – all parasitic, and – able to cause serious human diseases, such as
malaria caused by Plasmodium.
Protozoans
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Figure 15.22e
Red blood cell
Apical complex
TEM
An apicomplexan
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• Another apicomplexan is Toxoplasma, – occurring in the digestive tracts of millions of people
in the United States but – held in check by the immune system.
• A woman newly infected with Toxoplasma during pregnancy can pass the parasite to her unborn child, who may suffer nervous system damage.
Protozoans
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• Ciliates are protozoans that – are named for their use of hair-like structures called
cilia to move and sweep food into their mouths, – are mostly free-living (nonparasitic), such as the
freshwater ciliate Paramecium, and – include heterotrophs and mixotrophs.
Protozoans
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Figure 15.22f
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A ciliate
Cilia
Cell “mouth”
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Slime Molds
• Slime molds – resemble fungi in appearance and lifestyle due to
convergence, but – are more closely related to amoebas.
• The two main groups of these protists are – plasmodial slime molds and – cellular slime molds.
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• Plasmodial slime molds – are named for the feeding stage in their life cycle,
an amoeboid mass called a plasmodium, – are decomposers on forest floors, and – can be large.
Slime Molds
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Figure 15.23
A plasmodial slime mold
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• Cellular slime molds have an interesting and complex life cycle of successive stages: – a feeding stage of solitary amoeboid cells, – a swarming stage as a slug-like colony that can
move and function as a single unit, and – a stage during which they generate a stalk-like
multicellular reproductive structure.
Slime Molds
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Figure 15.24
Life stages of a cellular slime mold
Slug-like colony
Amoeboid cells
Reproductive structure
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Unicellular and Colonial Algae
• Algae are – photosynthetic protists whose chloroplasts support
food chains in – freshwater and – marine ecosystems.
• Many unicellular algae are components of plankton, the communities of mostly microscopic organisms that drift or swim weakly in aquatic environments.
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• Unicellular algae include – dinoflagellates, with
– two beating flagella and – external plates made of cellulose,
– diatoms, with glassy cell walls containing silica, and
– green algae.
Unicellular and Colonial Algae
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• Green algae are – unicellular in most freshwater lakes and ponds, – sometimes flagellated, such as Chlamydomonas,
and – sometimes colonial, forming a hollow ball of
flagellated cells as seen in Volvox.
Unicellular and Colonial Algae
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Figure 15.25
(a) A dinoflagellate, with its wall of protective plates
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(b) A sample of diverse diatoms, which have glassy walls
(c) Chlamydomonas, a unicellular green alga with a pair of flagella
(d) Volvox, a colonial green alga
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Figure 15.25a
(a) A dinoflagellate, with its wall of protective plates
SEM
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Figure 15.25b
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(b) A sample of diverse diatoms, which have glassy walls
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Figure 15.25c
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(c) Chlamydomonas, a unicellular green alga with a pair of flagella
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Figure 15.25d
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(d) Volvox, a colonial green alga
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Seaweeds
• Seaweeds – are large, multicellular marine algae, – grow on or near rocky shores, – are only similar to plants because of convergent
evolution, – are most closely related to unicellular algae, and – are often edible.
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• Seaweeds are classified into three different groups, based partly on the types of pigments present in their chloroplasts: 1. green algae, 2. red algae, and 3. brown algae (including kelp).
Seaweeds
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Figure 15.26
Brown algaeRed algaeGreen algae
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Evolution Connection: The Origin of Multicellular Life• Multicellular organisms have
– specialized cells that are dependent on each other and perform different functions, such as
– feeding, – waste disposal, – gas exchange, and – protection.
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• Colonial protists likely formed the evolutionary links between – unicellular and – multicellular organisms.
• The colonial green alga Volvox demonstrates one level of specialization and cooperation.
Evolution Connection: The Origin of Multicellular Life
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Figure 15.27-1
Unicellular protist
An ancestral colony may have formed when a cell divided and remained attached to its offspring.
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Figure 15.27-2
Unicellular protist
Locomotor cells
Food- synthesizing cells
Cells in the colony may have become specialized and interdependent.
An ancestral colony may have formed when a cell divided and remained attached to its offspring.
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Figure 15.27-3
Unicellular protist
Locomotor cells
Food- synthesizing cells Somatic
cells
Gamete
Additional specialization may have led to sex cells (gametes) and nonreproductive cells (somatic cells).
Cells in the colony may have become specialized and interdependent.
An ancestral colony may have formed when a cell divided and remained attached to its offspring.
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Figure 15.UN01
Bacteria
Archaea
Prokaryotes
Eukarya
Protists
Plants
Fungi
Animals
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Figure 15.UN02
Bacteria
Archaea
Prokaryotes
Eukarya
Protists
Plants
Fungi
Animals
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Figure 15.UN03
Major episode Millions of years ago
All major animal phyla establishedPlants and fungi colonize land
Origin of Earth
First multicellular organismsOldest eukaryotic fossilsAccumulation of O2 in atmosphereOldest prokaryotic fossils
5005301,2001,8002,4003,5004,600
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Figure 15.UN04Inorganic compounds
Abiotic synthesis of organic monomers
Abiotic synthesis of polymers
Formation of pre-cells
Self-replicating molecules
Membrane-enclosed compartment
Complementary chain
Polymer
Organic monomers2
1
3
4
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Figure 15.UN05
Spherical Rod-shaped Spiral
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Figure 15.UN06
Nutritional Mode Energy Source Carbon Source
Photoautotroph
Chemoautotroph
Photoheterotroph
Chemoheterotroph
Sunlight
Inorganic chemicals
Sunlight
Organic compounds
CO2
Organic compounds
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Figure 15.UN07
Bacteria
Archaea
Prokaryotes
Eukarya
Protists
Plants
Fungi
Animals
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Figure 15.UN08
Bacteria
Archaea
Prokaryotes
Eukarya
Protists
Plants
Fungi
Animals