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Evidence of Evolution: Homology
Homology:
Similarity that exists in species descended from common ancestor
Homology can be recognized and studied at three interacting levels:
Genetic
Developmental
Structural
Evidence of Evolution: Genetic Homology
Genetic homology:
Similarity in DNA nucleotide sequences, RNA nucleotide sequences, or amino acid sequences
Cytochrome C: a protein used in Electron Transport Chain
Evidence of Evolution: Developmental Homology
Developmental homology:
Similarity in developmental structures or processes
Example:
Early chick, human, and cat embryos have tails and structures called pharyngeal pouches:
Product of similar development processes inherited from common ancestor
Evidence: Comparative Anatomy
Similarities in patterns of embryonic development suggest shared ancestry
Organisms with very different ways of life often share common anatomical structures due to descendance from common ancestors
Evidence of Evolution: Structural Homology
Structural homology:
Similarity in adult morphology
For example, most vertebrates have a common structural plan in the limb bones
Vestigial traits: evolution is not perfection
Evolution’s Internal Consistency
Multiple data sets support idea that species have descended, with modification, from a common ancestor
Internal consistency:
Observation that data from independent sources agree in supporting predictions made by a theory
Evidence for the evolution of cetaceans—whales and dolphins—illustrates idea of internal consistency:
The fossil record: cetaceans are identified by unique ear bones
Phylogeny of the fossil cetaceans:
Indicates a gradual transition between terrestrial and aquatic forms
Evolution’s Internal Consistency Continued
Relative dating and absolute dating both support order of species indicated in phylogeny
Phylogeny of living whales and dolphins:
Estimated from similarities and differences in DNA sequences
Indicate that hippos are closest relative to cetaceans
Vestigial hip and hindlimb bones are found in some adult whales and dolphin embryos
Domains of Life
Bacteria, Archaea, and Eukarya are the three largest branches on the tree of life.
All bacteria and archaea are prokaryotic and unicellular, and they help form the microbiome.
Fundamental differences between bacteria and archaea:
Bacteria have cell walls made of peptidoglycan.
Archaea have unique phospholipids in the cell membranes.
Bacteria and archaea have different ribosome and RNA polymerase structures.
Archaea are more closely related to Eukarya than to Bacteria.
Biological Impact of Bacteria
Ancient, diverse, abundant, and ubiquitous lineages:
Oldest fossils are of 3.5-billion-year-old bacteria:
Eukaryotes not in fossil record until 1.75 BY later
10,000 species named and described, but hundreds of thousands likely exist:
~1000 microbes in human large intestine
~700 microbes in human mouth
Bacteria and archaea are amazingly abundant
A mere teaspoon of soil contains billions of microbial cells.
A liter of seawater contains a community of microbes equivalent to that of a large human city.
Microbes living under the ocean may make up to 10% of the world’s total living biomass.
They are found in every possible environment.
They are very diverse, and we are still discovering entire new phyla.
Some are extremophiles
Extremophiles live in unusual environments.
there are bacteria that live at a pH less than 1.0, at temperatures of 0°C under the ice, and in water 5–10 times saltier than seawater.
Studying extremophiles may help us understand the origin of life, since life probably evolved in a high-temperature, anoxic environment.
Astrobiologists use extremophiles as model organisms in the search for extraterrestrial life.
Extremophiles are useful in certain commercial and research applications.
A genetic engineering challenge is to produce quantities of a desired protein
In DNA cloning a human gene that produces an important protein is isolated.
insulin
It is then inserted into a bacterial plasmid.
As the bacteria multiply, large amounts of the gene, and thus the protein, are produced.
Pathogenic bacteria
Come from several different lineages in the domain Bacteria.
Pathogens tend to affect tissues at entry points into the body.
The germ theory of disease
Based on Koch’s postulates (four criteria that had to be met to demonstrate that a specific microbe causes a certain disease).
infectious diseases are caused by microbes (microscopic organisms).
Infectious diseases are spread in three main ways:
From person to person
From bites of insects or animals
From ingesting contaminated food or water, or environmental exposure
The germ theory’s immediate impact was in improving sanitation, greatly reducing mortality due to infectious disease.
How have Bacteria and Archaeans diversified?
Genetic variation through gene transfer
Transformation—when bacteria or viruses naturally take up DNA from the environment
Transduction—when viruses pick up DNA from one prokaryotic cell and transfer it to another cell
Conjugation—when genetic information is transferred by direct cell-to-cell contact
Ways to identify bacteria
Size, shape, and motility
Ways to identify bacteria: Cell wall composition
Gram-positive bacteria have a cell wall with abundant peptidoglycan, which stains dark purple when exposed to a Gram stain.
Gram-negative bacteria have a cell wall with a thin layer of peptidoglycan surrounded by a phospholipid bilayer. They stain light pink.
Gram stain analysis can predict sensitivity to certain drugs.
Ways to identify bacteria: Metabolic diversity
Bacteria and archaea are astonishingly diverse in the ways they acquire energy to make ATP and the carbon compounds they can use as building blocks.
There are three ways to acquire energy to produce ATP:
Phototrophs use light energy to energize electrons, producing ATP by photophosphorylation (light reactions of photosynthesis).
Chemoorganotrophs oxidize organic molecules with high potential energy, such as sugars (cellular respiration, fermentation).
Chemolithotrophs oxidize inorganic molecules with high potential energy, such as ammonia or methane (usually via cellular respiration).
There are two ways to acquire carbon :
Autotrophs use carbon dioxide or methane to build their own carbon-containing compounds.
Heterotrophs acquire carbon-containing compounds from other organisms.
Overall, there are six major “feeding strategies” (the six possible combinations of three methods of acquiring energy and two methods of acquiring carbon).
Plants, animals, fungi, and other eukaryotes use only two strategies.
Bacteria and archaea use all six.
Basic chemistry required for photosynthesis, cellular respiration, and fermentation originated in these lineages
Evolution of variations on each of these processes allowed prokaryotes to diversify into millions of species that occupy diverse habitats
Ecological Diversity and Global Impacts
Bacteria and archaea produce extremely sophisticated enzymes :
As a result, they can live in extreme environments and use toxic compounds as food
The complex chemistry and abundance of bacteria and archaea make them potent forces for global change
Bacteria and archaea have altered the chemical composition of the oceans, the atmosphere, and terrestrial environments for billions of years.
The Oxygen Revolution
No free molecular oxygen existed for first 2.3 billion years of Earth’s history
Cyanobacteria:
lineage of photosynthetic bacteria
first to perform oxygenic photosynthesis
were responsible for changing Earth’s atmosphere to one with a high concentration of oxygen
Nitrogen Fixation and the N-cycle
All organisms require nitrogen (N) to synthesize proteins and nucleic acids
Molecular nitrogen (N2) is abundant in atmosphere
Plants cannot use molecular nitrogen (N2) directly
Plant growth is often limited by the availability of nitrogen
Must obtain N from ammonia (NH3) or nitrate (NO3-)
Nitrogen fixation—certain bacteria and archaea are the only organisms capable of converting N2 to NH3:
Nitrogen-fixing bacteria live in close association with plants (e.g., in root structures called nodules)
Ecological Diversity and Global Impacts
Other bacteria and archaea convert ammonia to nitrates and nitrites, resulting in a complex nitrogen cycle.
Nitrogen-fixing legumes can be used to restore N levels in badly degraded soils
Nitrate Pollution
Widespread use of NH3 fertilizers causes pollution:
When NH3 is added to soil, much of it is used by bacteria as food
These bacteria then release nitrite or nitrate as waste products
Nitrates cause pollution in aquatic environments:
In an aquatic ecosystem, nitrates can decrease oxygen content, causing anaerobic “dead zones” to develop
Fungi are the master traders and recyclers in terrestrial ecosystems
Some fungi release nutrients from dead plants and animals into the soil; others obtain nutrients and then transfer them directly to living plants and animals.
Because they recycle key elements such as carbon, nitrogen, and phosphorus and because they transfer key nutrients to plants and animals, fungi profoundly influence ecosystem productivity and biodiversity.
In terms of nutrient cycling, fungi make the world go around.
Economic and Ecological Impacts
~300 species of fungi cause human illness:
This incidence is low compared to other organisms
Their major destructive impact is on our food supply :
Rusts, smuts, mildews, wilts, and blights cause billions of dollars of crop losses each year
Saprophytic fungi are responsible for losses due to spoilage
Fungi have many positive impacts:
They are the source for many antibiotics (ex. Penicillin)
Mushrooms are eaten in many cultures
Yeast (Saccharomyces cerevisiae) is used to make bread, cheese, soy sauce, beer, wine, and other foods
Fungal enzymes improve characteristics of foods such as fruit juice, candy, and meat
Saprophytic Fungi Accelerate the Carbon Cycle on Land
Saprophytes are fungi that digest dead plant material
Fungi help cycle carbons through terrestrial systems
The carbon cycle on land has two basic components:
Fixation of carbon by land plants
Release of C O2 from cellular respiration
For many carbon atoms, saprophytic fungi connect the two components
Analyzing Morphological Traits
Fungi have very simple bodies
Two growth forms exist:
Single-celled forms—yeasts
Multicellular, filamentous forms—mycelia (singular: mycelium)
Some species adopt both forms
All mycelia are dynamic
They constantly grow in the direction of food sources and die back in areas where food is running out
The body shape of a fungus can change almost continuously throughout its life
The Nature of Hyphae
Hyphae—the long, narrow filaments of mycelium
Because mycelia are made of branching networks of very thin hyphae:
Fungi have the highest surface-area-to- volume ratio of all multicellular organisms
Nutrient absorption is extremely efficient
Prone to drying out:
Thus most abundant in moist environments
Reproductive spores are resistant to drying out
Spores can endure dry periods and then germinate
What Adaptations Make Fungi Such Effective Decomposers?
Given enough time, fungi can turn even the hardest, most massive trees into soft soils
Large surface area of a mycelium makes nutrient absorption exceptionally efficient
Saprophytic fungi can grow toward the dead tissues that supply their food
Fungi must digest their food before they can absorb it
Fungi thus perform extracellular digestion:
Digestion that takes place outside the organism
Simple compounds resulting from enzymatic action are absorbed by hyphae
The two most abundant organic molecules on Earth are digested by fungi:
Lignin—in plant secondary cell walls
Cellulose—in plant primary and secondary cell walls
Fungi are much more closely related to animals than to land plants
Fungal infections in humans are more difficult to treat than bacterial infections:
Recent shared ancestry results in similar cellular and molecular structures
Drugs that disrupt fungal physiology are likely to damage humans
Key traits linking animals and fungi:
D N A sequence data
Both animals and fungi synthesize chitin
Flagella in chytrid spores and gametes are similar to animal flagella
Animals and fungi store glucose as the polysaccharide glycogen
Fundamental features of eukaryotes
The nuclear envelope is a synapomorphy that defines the domain Eukarya.
Most eukaryotic cells are larger than prokaryotes, have many organelles, and have an extensive system of structural proteins called the cytoskeleton.
Multicellularity has evolved multiple times in eukaryotes.
Eukaryotes reproduce either asexually via mitosis or sexually via meiosis.
Protists are all the eukaryotes that are not fungi, green plants, or animals.
a collection of lineages which surrounded by water most of the time.
Three important evolutionary innovations in protists
evolution of the nuclear envelope/membrane
origin of the mitochondrion
origin of chloroplasts
Origin of the Nuclear Envelope
The evolution of the nuclear envelope was advantageous because it separated transcription and translation.
in bacteria and archaea, transcription and translation occur together
in eukaryotes RNA transcripts are processed inside the nucleus but translated outside the nucleus
With a simple nuclear envelope in place, alternative splicing and other forms of RNA processing could occur.
This important morphological innovation gave the early eukaryotes a novel way to control gene expression.
The endosymbiotic theory
Lynn Margulis proposed in the 1970s that mitochondria evolved from an aerobic bacterium that was engulfed by an anaerobic eukaryotic cell
Mutually beneficial symbiosis:
The host supplied the bacterium with protection and carbon compounds
the bacterium produced much more ATP than the host could produce on its own.
Many lines of evidence support Endosymbiotic Theory
Mitochondria are similar in size to alpha-proteobacteria. They:
divide independently of the host cell, and by fission, as bacteria do.
have their own ribosomes (similar to bacterial ribosomes) and synthesize their own proteins.
have double membranes, as would be expected if they were engulfed by another cell.
have their own chromosomes, which are circular and similar to bacterial chromosomes.
The most conclusive evidence is that mitochondrial genes are very closely related to the genes from alpha-proteobacteria
Endosymbiosis and the origin of the chloroplasts
All photosynthetic protists have chloroplasts.
None of the basic machinery required for photosynthesis evolved in eukaryotes.
they likely “captured” it via endosymbiosis
The eukaryotic chloroplast may have originated when a protist engulfed a cyanobacterium.
Once inside the protist,
the photosynthetic bacterium provided its eukaryotic host with oxygen and glucose
the host provided the bacterium protection and access to light
Which eukaryote originally obtained a photosynthetic organelle?
Because all species in the Plantae have chloroplasts with two membranes, biologists infer that the original, or primary, endosymbiosis occurred in these species’ common ancestor.
That ancestor eventually gave rise to all subgroups in the Plantae lineage—the glaucophyte algae, red algae, and green plants (green algae and land plants).
Evidence for endosymbiosis and the origin of the chloroplasts
Chloroplasts have the same list of bacteria-like characteristics presented earlier for mitochondria.
They contain circular DNA containing genes extremely similar to genes found in various species of cyanobacteria.
Some algae have a photosynthetic organelle with an outer layer containing the same peptidoglycan found in cyanobacteria.
There are many examples of endosymbiotic cyanobacteria living inside cells of protists or animals today.
Secondary Endosymbiosis Leads to Organelles with Four Membranes
In Excavata, Rhizaria, Alveolata, and Stramenopila, the chloroplast is surrounded by more than two membranes—usually four.
Researchers hypothesize that the ancestors of these groups acquired their chloroplasts by ingesting photosynthetic protists that already had chloroplasts.
Secondary endosymbiosis: occurs when an organism engulfs a photosynthetic eukaryotic cell and retains the chloroplasts as intracellular symbionts.
Similarities Between Green Algae and Land Plants
Of the green algal groups, three most similar to land plants:
Based on DNA sequence analysis:
Zygnematophyceae (conjugating algae)
Coleochaetophyceae (coleochaetes)
Charophyceae (stoneworts)
Largely multicellular and live in freshwater:
Hypothesis: Land plants evolved from green algae that lived in freshwater habitats
How Do Biologists Study Green Algae and Land Plants?
To understand diversification:
Compare morphological traits
Analyze the fossil record
Estimate phylogenetic trees
Plants: Life on Land
The first-known green algae lived between 700 and 725 million years ago (mya), when oxygen levels began to rise.
Green algae (protists) live surrounded by water
water, minerals
Support
reproduction
Plants: Life on Land (475 mya)
Land plants are monophyletic :
There was only one successful transition from freshwater environments to land
To adapt to life on land, plants evolved features that allow them to
resist drying out (absorb and retain water)
absorb nutrients
stand upright without outside support
reproduction not dependent on water