BIS2C_2020. Lecture 8. Phylogenetic Diversity of Microbes

Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2020
BIS2C
Biodiversity & the Tree of Life
Spring 2020
Lecture 8:
Phylogenetic Diversity of Microbes
Prof. Jonathan Eisen

Anna’s Hummingbird in Davis, CA April 8, 2020

Learning Goals
1. Understand what type of data has been used to infer
the overall major structures in the Tree of Life
2. Understand the great plate count anomaly and how it
has impacted our view of the phylogenetic diversity of
life
3. Understand and be able to give examples of key
features present in key ancestors of major branches
in the tree of life (e.g., LUCA, BCA, ECA)

!7: The Domains of Life
!8: Phylogenetic Diversity of Microbes
!9: Functional Diversity of Microbes
Lecture 8 Context

Lecture 8 Outline
• Background
• Phylogenetic Diversity of Life
• Features and Innovations of Major Groups
• Bonus Tours

Background: Lab Connections
• Lab 2 - Intro to Bacteria and Archaea
• Lab 2 - Station A - classification
• Lab 3 - Intro to microbial eukaryotes
• Lab 2 - Great plate count anomaly

Background: Review of Lecture 7

Cell Structure Observations
L7: Prokaryotes vs. Eukaryotes

EukaryotesProkaryotes
No Nucleus Nucleus
Two Main Kinds of Structures
What Other Differences Are There?

No Nucleus Nucleus
Mitochondria
Endoplasmic
Reticulum
Diploid
MeiosisBinary
Fission
Mitosis
Golgi
Linear DNACircular DNA
Haploid
Chromatin

Prokaryotic
Eukaryotic
P1 P2 P3 E1 E2 E3P4
Interpreted as
implying this tree
because this would
require only one event
for the nucleus origin

P1 P2 P3 E1 E2 E3P4
Prokaryotic
Eukaryotic
Interpreted As Implying This Tree Because That Would
Require Only One Event For Nucleus Origin
But this is a mistake
since this tree, also
only requires one
event for origin of
the nucleus

Prokaryotic
Eukaryotic
P1 P2 P3 E1 E2 E3P4
So no reason to
assume this is the
correct tree, even
though many did

L7: Woese rRNA Tree of Life

rRNA rRNArRNA
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
R ACUCCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
F ACUCCAGGUAUCGAUCG
C ACCCCAGCUCUCGCUCG
W ACCCCAGCUCUGGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
C ACCCCAGCUCUCGCUCG
L7: Woese rRNA Tree of Life

P1 P2 P3 E1 E2 E3P4 P5 P6 P7
Prokaryotic
Eukaryotic
L7: Rooted Woese rRNA Tree of Life

B1 B2 B3 E1 E2 E3B4 A1 A2 A3
Prokaryotic
Eukaryotic
L7: The Three Domain Tree of Life
EukaryaBacteria Archaea
Three Domains
Re-labelled
individual taxa

L7: Loki’s Castle
• Loki’s Castle

B1 B2 B3 E1 E2 E3B4 A1 A2 Loki
Prokaryotic
Eukaryotic
EukaryaBacteria Archaea
Phylogenetic Trees
Using Genomes from
Lokiarchaea Look Like
This
L7: Alternative to Three Domain Tree of Life

B1 B2 B3 E1 E2 E3B4 A1 A2 Asgard
Prokaryotic
Eukaryotic
Bacteria
L7: Two Domain Tree of Life
Two Domain Tree of Life
Archaea
Eukarya
“Prokaryotic
Archaea”

Prokaryotic
Eukaryotic
Bacteria
L7: Three Domain Tree of Life
Three Domain Tree of Life
Archaea Eukarya
Note Position of Asgard Archaea
New Slide

• Method developments
!Same data, with different methods,
sometimes gives different results
!Some methods are more prone to artifacts
and errors than others
• Differences between genes
• Better taxonomic sampling
!Can improve resolution in trees
!Can help get around some method problems
Reasons for Changes in the Tree of Life
L7: Two Domain Tree of Life

Lecture 8 Outline
• Background and Context
• Bonus tours

Prokaryotic
Eukaryotic
Bacteria
Archaea
Eukarya
“Prokaryotic
Archaea”

Key Lesson #1
• The overall structure of the tree of life has
been determined by analysis of molecular
sequence data
• Molecular data particularly important for
placing most microbes into the tree of life

Woese rRNA Tree of Life

rRNA rRNArRNA
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
R ACUCCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
F ACUCCAGGUAUCGAUCG
C ACCCCAGCUCUCGCUCG
W ACCCCAGCUCUGGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
C ACCCCAGCUCUCGCUCG

genome genomegenome
ACUGCACCUAU
CGUUCGACUGCACC
UAUCGUUCGACUGC
ACCUAUCGUUCGAC
UGCACCUAUCGUUC
GACCUAUCGUUCGA
CUGCACCUAUCGUU
CGACCUAUCGUUCG
ACUGCACCUAUCGU
UCG
ACUGCACCUAU
CGUUCGACUGCACCU
AUCGUUCGACUGCAC
CUAUCGUUCGACUGC
ACCUAUCGUUCGACC
UAUCGUUCGACUGCA
CCUAUCGUUCGACCU
AUCGUUCGACUGCAC
ACUGCACCUAU
CGUUCGACUGCACCU
AUCGUUCGACUGCAC
CUAUCGUUCGACUGC
ACCUAUCGUUCGACC
UAUCGUUCGACUGCA
CCUAUCGUUCGACCU
AUCGUUCGACUGCAC
CUAUCGUUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
R ACUCCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
F ACUCCAGGUAUCGAUCG
C ACCCCAGCUCUCGCUCG
W ACCCCAGCUCUGGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
C ACCCCAGCUCUCGCUCG

B1
B2
B3
E1
E2
E3
B4
A1
A2
Asgard
Bacteria
Archaea
Eukarya
“Prokaryotic
Archaea”

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Two Domain Tree of
Life with Some Major
Subbranches Shown

Key Lesson #2
• Most microbes were only included in the tree
of life if they could be grown in the lab

Great Plate Count Anomaly

<<<<
Culturing Observation
CountCount

<<<<
Culturing Observation
CountCount
http://www.google.com/url?
sa=i&rct=j&q=&esrc=s&source=images&cd
=&docid=rLu5sL207WlE1M&tbnid=CRLQYP
7d9d_TcM:&ved=0CAUQjRw&url=http%3A%
2F%2Fwww.biol.unt.edu%2F~jajohnson%2F
DNA_sequencing_process&ei=hFu7U_TyCtO
qsQSu9YGwBg&psig=AFQjCNG-8EBdEljE7-
yHFG2KPuBZt8kIPw&ust=140487395121142
4
DNA
Lab 2:
“Culture Independent
DNA Studies”

Taxa Characters
S ACUGCACCUAUCGUUCG
R ACUCCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
F ACUCCAGGUAUCGAUCG
C ACCCCAGCUCUCGCUCG
W ACCCCAGCUCUGGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
C ACCCCAGCUCUCGCUCG
Environmental Sampling
genome genomegenome
ACUGCACCUAU
CGUUCGACUGCACC
UAUCGUUCGACUGC
ACCUAUCGUUCGAC
UGCACCUAUCGUUC
GACCUAUCGUUCGA
CUGCACCUAUCGUU
CGACCUAUCGUUCG
ACUGCACCUAUCGU
UCG
ACUGCACCUAU
CGUUCGACUGCACCU
AUCGUUCGACUGCAC
CUAUCGUUCGACUGC
ACCUAUCGUUCGACC
UAUCGUUCGACUGCA
CCUAUCGUUCGACCU
AUCGUUCGACUGCAC
ACUGCACCUAU
CGUUCGACUGCACCU
AUCGUUCGACUGCAC
CUAUCGUUCGACUGC
ACCUAUCGUUCGACC
UAUCGUUCGACUGCA
CCUAUCGUUCGACCU
AUCGUUCGACUGCAC
CUAUCGUUCG

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Most of the
phylogenetic diversity
of bacteria only known
through DNA from
environmental
samples

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Most of the
through DNA from
environmental
samples
Same is true for
Archaea

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Most of the
through DNA from
environmental
samples
Same is true for
Archaea
Also true for the
microbial lineages in
eukaryotes

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Most of the
through DNA from
environmental
samples
Same is true for
Archaea
Also true for the
microbial lineages in
eukaryotes
No way to cover all
this diversity in this
class

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Focus on this reduced
subset of the full

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Plants: L14-21
Fungi L 22-23
Animals L 25-36
Choanos: L 24
Everything Else L 8-13
NOOP
Not Opisthokonts or Plants
subset of the full

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
L8: Features and
Innovations of Major
Groups
L9: Functional Diversity
L10: Parasites and
Pathogens
L11: Viruses and Gene
Transfer
L12: Mutualisms and
Microbiomes
L13: Endosymbioses and
Organelles
Everything Else L 8-13
NOOP
Not Opisthokonts or Plants
subset of the full

Lecture 8 Outline
• Bonus Tours

Lecture 8 Outline
!LUCA
• Bonus Tours

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
LUCA
LUCA
Last
Universal
Common
Ancestor

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
LUCA
LUCA
Last
Universal
Common
Ancestor
Can infer
traits using
character
state
reconstruction
methods

• Made up of cells
• Use of DNA as a genetic material
• Use of ACTG in DNA
• Use of ACUG in RNA
• Three letter genetic code
• Central dogma (DNA » RNA » protein)
• Use water as a solvent
• Lipoprotein cell envelope
• 20 core amino acids in proteins
• Live on Earth
• Ribosome for translation
• RNA polymerase proteins
• Acquires energy from environment
• Store energy in chemicals
Universal Traits

• Made up of cells
• Use of DNA as a genetic material
• Use of ACTG in DNA
• Use of ACUG in RNA
• Three letter genetic code
• Central dogma (DNA » RNA » protein)
• Use water as a solvent
• Lipoprotein cell envelope
• 20 core amino acids in proteins
• Live on Earth
• Ribosome for translation
• RNA polymerase proteins
• Acquires energy from environment
• Store energy in chemicals
Universal Traits
Infer that
LUCA had all
of these

Prokaryotes vs. Eukaryotes
No Nucleus Nucleus

Prokaryotes vs. Eukaryotes
No Nucleus Nucleus
Mitochondria
Endoplasmic
Reticulum
Diploid
MeiosisBinary
Fission
Mitosis
Golgi
Linear DNACircular DNA
Haploid
Chromatin

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Circular DNA
Linear DNA

Traits of LUCA = Last Universal Common Ancestor
• For characters that differ between prokaryotes
and eukaryotes, LUCA likely had the
character / character state found in
prokaryotes
!Circular DNA genome with plasmids
!No nucleus or membrane bound organelles
!Operons
!Binary fission reproduction
!Haploid

Lecture 8 Outline
!LUCA
!BCA
• Bonus Tours

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
BCA
BCA
Bacterial
Common
Ancestor

BCA = Bacterial Common Ancestor
• Likely inherited most of the traits inferred to
be in LUCA
• A few other key innovations, we are just going
to discuss one today

Peptidoglycan
• Unique to bacteria
• Polymer that provides a
firm, mesh like structure
around the cell
• Key component of cell
wall
• Is the target for many
antibiotics

Peptidoglycan
• Unique to bacteria
• Polymer that provides a
firm, mesh like structure
around the cell
• Key component of cell
wall
• Is the target for many
antibiotics
• Why?

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
PG
Peptidoglycan
is a key
innovation of
bacteria

Peptidoglycan variation in bacteria
• Two major ways it is
used in bacteria
• Gram-positive bacteria -
thick peptidoglycan layer
outside 1 cell membrane
• Gram-negative bacteria
- thin peptidoglycan
layer outside between
two cell membranes
• This has many impacts
on biology, including
sensitivity to antibiotics

Lecture 8 Outline
!LUCA
!BCA
!ECA
• Bonus Tours

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
ECA
Eukaryote
Common
Ancestor

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
ECA
Eukaryote
Common
Ancestor
Tons of
innovations
thought to have
happened here

Many Other Innovations Associated w/ Origin of Euks
• Cell Structure
!Cytoskeleton
!Phagocytosis
!Diverse organelles and compartments
• Molecular Processes
!Mitosis (vs. binary fission)
!Sexual reproduction (vs. gene transfer)
" Meiosis
" Haploidy vs. Diploidy
" Fertilization
!Linear chromosomes & telomeres
!Chromosomes packaged into chromatin
!Decoupling of transcription and translation
!Introns and splicing

Lecture 8 Outline
!LUCA
!BCA
!ECA
!Asgard Archaea and Eukaryotes

Lokiarchaea named after Loki’s Castle Vent Site

Found in Asgard Archaea
• Cell Structure
!Cytoskeleton
!Phagocytosis
!Diverse organelles and compartments
• Molecular Processes
!Mitosis (vs. binary fission)
!Sexual reproduction (vs. gene transfer)
" Meiosis
" Haploidy vs. Diploidy
" Fertilization
!Linear chromosomes & telomeres
!Chromosomes packaged into chromatin
!Decoupling of transcription and translation
!Introns and splicing

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Some features
previously
thought to be
“Eukaryotic”
innovations
were likely
present in the
Eukaryote-
Asgard
ancestor
Cytoskeleton
Chromatin

Tour of Major Clades
• You Do Not Need to Know These Details
• Really

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Bacteria
Tour

Spirochetes
• Gram-negative
• Motile
• Chemoheterotrophic
• Unique rotating, axial
filaments (modified flagella)
• Many are pathogens:
!Syphilis
!Lyme disease
• Others free-living

Chlamydias
• Gram-negative
• Cocci or rod-shaped
• Extremely small
• Live only as parasites
inside cells of
eukaryotes & cause
various diseases
!Trachoma
!Multiple sexually
transmitted diseases
!Pneumonia
C. trachomatis

Cyanobacteria
• Photolithoautotrophic
• Contain internal
membrane system
for photosynthesis
• Chloroplasts are
derived from
endosymbiotic
cyanobacteria
• Colonies can
differentiate into
vegetative cells,
spores, & heterocysts

Proteobacteria
• Gram-negative
• Escherichia coli: model
organism and human gut
commensal and
pathogen
• Mitochondria evolved
from this group
• Includes many human
and animal pathogens:
plague, cholera, typhoid

Low-GC Gram Positives (Firmicutes)
• Low G+C/A+T ratio in DNA
• Some produce endospores
which are resistant “seeds”
that germinate when
conditions are good
• Many agents of diseases
(e.g., anthrax, MRSA,
Streptococcus, botulism,
tetanus)
• Many of agricultural and
industrial use (e.g., Lactic
acid bacteria)
• Some (Mycoplasmas) have
no cell wall and are
extremely small
Mycoplasmas

High-GC Gram Positives (Actinobacteria)
• High G+C/A+T ratio in DNA
• Elaborate branching
• Some reproduce by forming
chains of spores at tips of
filaments
• Most antibiotics are from
this group
• Causative agents of many
diseases such as
tuberculosis and leprosy
• Many originally misclassified
as fungi

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Prokaryotic
Archaea
Tour

Crenarchaeota
• Most are both
thermophilic (heat
loving) & acidophilic
(acid-loving)
• Sulfolobus lives in
hot sulfur springs
(70–75°C, pH 2-3)
• One species of
Ferroplasma lives at
pH near 0

Euryarchaeota: Methanogens
• Only known methanogens
(produce methane (CH4)
by reducing CO2 ) - a form
of chemoautotrophy
• Methanogens release 2
billion tons of methane per
year
• Many live in the guts of
grazing mammals
• Many such as
Methanopyrus live in deep-
ocean hydrothermal vents

Euryarchaeota: Halophiles (Salt lovers)
• Pink carotenoid
pigments – very visible
• Have been found at
pH up to 11.5.
• Unusual adaptations
to high salt,
desiccation
• Many have
bacteriorhodopsin
which uses energy of
light to synthesize ATP
(photoheterotrophs)

Opisthokonts
Amoebozoans
Excavates
Plantae
Rhizaria
Stramenopiles
Alveolates
Eukarya
Tour

Plantae
The Plantae consist of
several clades; all
chloroplasts trace back to a
single incidence of
endosymbiosis.

Plantae: Glaucophytes
• Unicellular, freshwater
organisms
• The chloroplast retains a bit
of peptidoglycan between the
inner and outer membrane.

Plantae: Red Algae
• Most red algae are marine and
multicellular.
• Red pigment is phycoerythrin.
•Many reproduce with spores
Motile spores from
Purpureoﬁlum
Audouinella paciﬁca
Spyridia

Plantae: Chlorophytes
• Sister group to charophytes and land
plants.
• Synapomorphies include chlorophyll a
and b, and starch as a storage product.
• >17,000 species; marine, freshwater,
and terrestrial. Unicellular to large
multicellular forms.
Movement in the green alga
Volvox
Micrasterias

Plantae: Charophytes and Land Plants
STAY
TUNED

Opisthokonts

•Choanoflagellates are sister to the
animals.
•Some are colonial and resemble a type
of cell found in sponges.
The choanoﬂagellate Salpingoeca sp. feeding
Opisthokonts: Choanoflagellates

Opisthokonts: Fungi and Animals
STAY
TUNED

Alveolates
Alveolates
Have alveoli or
sacs beneath
surface of
plasma
membrane.
All are unicellular;
many are
photosynthetic.

•Most are marine and are important photoautotrophic
primary producers
•Mixture of pigments give them a golden brown color.
•Have two flagella, one in an equatorial groove, the other
in a longitudinal groove.
Alveolates: Dinoflagellates
Certium
tenue
Coral symbiont

Alveolates: Apicomplexans
• All parasitic
• Have a mass of organelles at one tip—the
apical complex that help the parasite enter
the host’s cells.
Apical complex • Plasmodium falciparum-
Malaria kills 700,000-2,000,000
people per year—75% of them
are African children

Alveolates: Ciliates
Movement in a ciliate from the gut of a termite
• All have numerous cilia, the structure is
identical to flagella.
• Most are heterotrophic; very diverse group.
• Have complex body forms and two types of
nuclei.

Stramenopiles
Stramenopiles
Two flagella, with rows of tubular hairs
on the longer one.

•All are multicellular; some get very large (e.g., giant
kelp).
•The carotenoid fucoxanthin imparts the brown color.
•Almost exclusively marine.
Stramenopiles: Brown Algae
A community of brown algae: The marine kelp forest

Stramenopiles: Diatoms
A colony of the diatom, Bacillaria
paradoxa
•Unicellular, but many associate in filaments.
•Have carotenoids and appear yellow or
brown.
•Excellent fossil record
•Most are photoautotrophic
•Responsible for 20% of all carbon fixation.
•Oil, gas source

Stramenopiles: Oomcyetes
Phytophthora
Potato Late Blight
• Non-photosynthetic.
• Are absorptive heterotrophs
• Once were classed as fungi, but are
unrelated.
Sudden Oak Death

Rhizaria
Rhizaria
Unicellular, aquatic, with long, thin pseudopods.

Rhizaria: Cercozoans
Some cercozoans are aquatic, others live
in soil.
They have diverse forms and habitats.
One group has chloroplasts derived from a
green alga by secondary endosymbiosis.
Euglyphid
Chlorarachnion reptans

Rhizaria: Foraminiferans
Sand beaches in the tropics
• Secrete shells of calcium carbonate.
• Discarded shells make up limestone.
• Create some beach sands
• Used to date & characterize sedimentary
rocks.
• Some live as plankton, others at sea bottom.
• Thread-like, branched pseudopods extend
through pores in the shell and form a sticky net
that captures smaller plankton.

Rhizaria: Radiolarians
• Have thin, stiff pseudopods reinforced
by microtubules.
• The pseudopods increase surface
area for exchange of materials; and
help the cell float.
• Exclusively marine, most secrete
glassy endoskeletons, many with
elaborate designs.

Excavates

Excavates: Diplomonads and Parabisalids
• Unicellular
• Lack mitochondria and most are
anaerobic. This is a derived condition
• Giardia lamblia - a diplomonad - is a
human parasite
• Trichomonas vaginalis - parabasalid - STD

Excavates: Heteroloboseans
• Amoeboid body form.
• Naegleria can enter humans and
cause a fatal nervous system
disease - “brain eating”
• Some can transform between
amoeboid and flagellated stages.

Excavates: Euglenids
• Have flagella.
• Some are
photosynthetic,
some always
heterotrophic, and
some can switch.
Movement in the euglenoid Eutreptia

Excavates: Kinetoplastids
• Unicellular parasites with two flagella and a
single mitochondrion.
• Mitochondrion contains a kinetoplast -
structure with multiple, circular DNA
molecules
• Includes trypanosomes and agents of
chagas, sleeping sickness, Leishmaniasis
Trypanosoma sp.
mixed with blood cells

Amoebozoans
Amoebozoans
Lobe-shaped pseudopods are used for
locomotion.

• Not colonial; live as single cells
• Some secrete shells or glue sand
grains together to form a casing.
• Many pathogens
Amoebozoans: Loboseans

Entamoeba histolytica
http://www.npr.org/blogs/health/2014/04/09/300991364/gut-eating-amoeba-
caught-on-ﬁlm

Amoebozoans: Plasmodial Slime Molds
• Individual motile cells can form single,
multinucleate cell (plasmodium)
• Ingest food by endocytosis
• Form spores on stalks called fruiting
bodies.
• Found in cool, moist habitats

Amoebozoans: Cellular Slime Molds
• Life cycle consists of individual motile cells that
ingest food by endocytosis
• This is followed by the formation of single,
multicellular fruiting structure
• Each cell retains its own plasma membrane and
individuality
Karyog

End of The Tours
• End of the Tours

!7: The Domains of Life
!8: Phylogenetic Diversity of Microbes
!9: Functional Diversity of Microbes
Lecture 8 Context
• See you Monday for Functional Diversity of
Microbes

BIS2C_2020. Lecture 8. Phylogenetic Diversity of Microbes

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