The document discusses a lecture on microbial diversity. It notes that the tree of life is mostly microbial, diverse methods exist to study microbial diversity, and most microbial diversity remains poorly characterized. Sequencing methods like rRNA and metagenomic sequencing have improved understanding of microbial phylogeny but much diversity remains unknown.
1. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Lecture 9:
Microbial Diversity
BIS 002C
Biodiversity & the Tree of Life
Spring 2016
Prof. Jonathan Eisen
1
2. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Where we are going and where we have been
• Previous Lecture:
!8: The Tree of Life II
• Current Lecture:
!9: Microbial Diversity
• Next Lecture:
!10: The Not Quite a Tree Tree of Life
2
3. Lectures 9-13 Microbial Diversity
• The Tree of Life is mostly microbial
• Diverse methods are available for studying microbial
diversity
• Most of the diversity of microbial life is poorly
characterized
• The Tree of Life is not actually a tree
• The biological diversity (form, function, etc) seen in
microbes is immense
• Microbes run (kind of) the planet
• Microbial interactions (with each other and non-
microbes) also help run the planet
3
4. Lectures 9
• The Tree of Life is mostly microbial
• Diverse methods are available for studying microbial
diversity
• Most of the diversity of microbial life is poorly
characterized
• The Tree of Life is not actually a tree
• The biological diversity (form, function, etc) seen in
microbes is immense
• Microbes run (kind of) the planet
• Microbial interactions (with each other and non-
microbes) also help run the planet
4
5. Lectures 10
• The Tree of Life is mostly microbial
• Diverse methods are available for studying microbial
diversity
• Most of the diversity of microbial life is poorly
characterized
• The Tree of Life is not actually a tree
• The biological diversity (form, function, etc) seen in
microbes is immense
• Microbes run (kind of) the planet
• Microbial interactions (with each other and non-
microbes) also help run the planet
5
6. Lecture 11
• The Tree of Life is mostly microbial
• Diverse methods are available for studying microbial
diversity
• Most of the diversity of microbial life is poorly
characterized
• The Tree of Life is not actually a tree
• The biological diversity (form, function, etc) seen in
microbes is immense
• Microbes run (kind of) the planet
• Microbial interactions (with each other and non-
microbes) also help run the planet
6
7. Lectures 12-13
• The Tree of Life is mostly microbial
• Diverse methods are available for studying microbial
diversity
• Most of the diversity of microbial life is poorly
characterized
• The Tree of Life is not actually a tree
• The biological diversity (form, function, etc) seen in
microbes is immense
• Microbes run (kind of) the planet
• Microbial interactions (with each other and non-
microbes) also help run the planet
7
8. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Lecture 9: Microbial Diversity
• The Tree of Life is mostly microbial
• Diverse methods are available for studying
microbial diversity
• Most of the diversity of microbial life is
poorly characterized
8
9. Unrooted Tree of Life (from ~ 2004)
9
adapted from Baldauf, et al., in Assembling the Tree of Life, 2004
10. 10adapted from Baldauf, et al., in Assembling the Tree of Life, 2004
P
Plants
Two Weeks
Unrooted Tree of Life (from ~ 2004)
11. 11adapted from Baldauf, et al., in Assembling the Tree of Life, 2004
F
Fungi
One Week
P
Unrooted Tree of Life (from ~ 2004)
12. 12adapted from Baldauf, et al., in Assembling the Tree of Life, 2004
A
Animals
Two Weeks
P
Unrooted Tree of Life (from ~ 2004)
13. 13adapted from Baldauf, et al., in Assembling the Tree of Life, 2004
1.5 Weeks
Unrooted Tree of Life (from ~ 2004)
14. Unrooted Tree of Life
14adapted from Baldauf, et al., in Assembling the Tree of Life, 2004
1.5 Weeks
Mostly Microbes
15. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
The Bacteria and Archaea via Textbook v.10
15
16. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Eukaryotic Groups via Textbook v.10
16
17. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014
Eukaryote Groups - More Detail
1717
18. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Microbial Diversity
• The Tree of Life is mostly microbial
• Diverse methods are available for studying
microbial diversity
• Most of the diversity of microbial life is
poorly characterized
18
19. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016 20
Field Observations Are Important Tools
20. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
• Field studies of multicellular organisms are
of course common
• Show binoculars, butterfly nets, etc
• Field studies of microbes are also possible
but a bit more challenging
21
Field Observations Important in Microbial Studies
21. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
• More detail on some of these in Labs 2 and
3
22
22. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016 !23
Culturing Microbes
23. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Culturing
• More On This in Lectures 11-12
• Some in Labs 2-3
24
24. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
DNA Sequencing
• All cellular organisms have genomes made up
of DNA
• All cellular organisms transcribe DNA into
RNA and then translate RNA into protein
• Sequencing involves reading the string of
letters in DNA, RNA or protein
• Sequencing is usually done on DNA
• Sequencing gets cheaper and faster VERY
fast
• Sequencing is very useful is studying
microbial diversity
25
25. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016 26
Sequencing Has Gone Crazy
26. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
rRNA Sequencing for Phylogenetic Analysis
27
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
EukaryotesBacteria ?????Archaea
28. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Which of the following is NOT a reason that
analysis of rRNA is useful for inferring a Tree
of Life
A: rRNAs are universal homologies
B: rRNAs can be sequenced
C: rRNAs are transcribed from DNA
D: rRNAs have functional roles in ribosomes
E: rRNAs don't vary between species
29
29. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Which of the following is NOT a reason that
analysis of rRNA is useful for inferring a Tree
of Life
A: rRNAs are universal homologies
B: rRNAs can be sequenced
C: rRNAs are transcribed from DNA
D: rRNAs have functional roles in ribosomes
E: rRNAs don't vary between species
30
30. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Genome Sequencing Improves Phylogenetic Analysis
31
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
EukaryotesBacteria ?????Archaea
DNA DNADNA
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
31. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Genome Sequencing Has Many Other Uses
32
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
DNA DNADNA
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
32. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Genome Sequencing Has Many Other Uses
33
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
DNA DNADNA
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
Some Discussion of this in Lecture 10
33. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
2002 Most Genomes from a Few Groups
34
Figure from Barton, Eisen et al. “Evolution”, CSHL Press based on Baldauf et al Tree
34. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
2002-2007: TIGR Tree of Life Project
35
Figure from Barton, Eisen et al. “Evolution”, CSHL Press based on Baldauf et al Tree
Naomi
Ward
Karen
Nelson
35. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
2007-2014: Genomic Encyclopedia
36
Figure from Barton, Eisen et al. “Evolution”, CSHL Press based on Baldauf et al Tree
36. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
2007-2014: Genomic Encyclopedia
37
Figure from Barton, Eisen et al. “Evolution”, CSHL Press based on Baldauf et al Tree
BUT …
37. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Microbial Diversity
• The Tree of Life is mostly microbial
• Diverse methods are available for studying
microbial diversity
• Most of the diversity of microbial life is
poorly characterized
38
38. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Great Plate Count Anomaly
39
<<<<
Culturing Observation
CountCount
39. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Great Plate Count Anomaly
39
<<<<
Culturing Observation
CountCount
http://www.google.com/url?
sa=i&rct=j&q=&esrc=s&source=images&c
d=&docid=rLu5sL207WlE1M&tbnid=CRLQ
YP7d9d_TcM:&ved=0CAUQjRw&url=http
%3A%2F%2Fwww.biol.unt.edu
%2F~jajohnson
%2FDNA_sequencing_process&ei=hFu7U_
TyCtOqsQSu9YGwBg&psig=AFQjCNG-8EB
dEljE7-
yHFG2KPuBZt8kIPw&ust=14048739512114
24
DNA
40. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
rRNA Sequencing from Environmental Samples
40
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
EukaryotesBacteria ?????Archaea
41. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Metagenomics Improves Phylogenetic Resolution
41
metagenomics
ACUGC
ACCUAU
CGUUCG
ACUCC
AGCUAU
CGAUCG
ACCCC
AGCUCU
CGCUCG
Taxa Characters
S ACUGCACCUAUCGUUCG
R ACUCCACCUAUCGUUCG
E ACUCCAGCUAUCGAUCG
F ACUCCAGGUAUCGAUCG
C ACCCCAGCUCUCGCUCG
W ACCCCAGCUCUGGCUCG
EukaryotesBacteria Archaea
Jo
Handelsman
43. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016 43
Sequencing Has Gone Crazy
44. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Hug et al. 2016
• Dowloaded 10,000+ genomes
from various databases
(including many I generated)
• 1000+ new genomes
• Searched these genomes for
a set universal homologous
genes (ribosomal proteins)
(based on AMPHORA)
• Aligned the sequences of
these genes between species
• Maximum likelihood tree
44
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
Laura Hug
U. Waterloo
Jill Banfield
UC Berkeley
45. Hug et al 2016
!45
Hug et al. 2016 Tree of Life
92 Bacterial Phyla
25 Archaeal Phyla
5 Eukaryotic Supergroups
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
Laura Hug
U. Waterloo
Jill Banfield
UC Berkeley
46. !46
Hug et al 2016Hug et al. 2016 Bacteria
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
47. Taxa Covered in Textbook
!47
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
48. !48
Hug et al 2016Phyla Never Grown in the Lab
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
49. Hug et al 2016
!49
Hug et al. 2016 Archaea and Eukaryotes
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
50. Hug et al 2016
!50
Hug et al. 2016 Archaea Phyla Never Grown in the Lab
Hug et al. Nature Microbiology. A new view of the tree of life.
http://dx.doi.org/10.1038/nmicrobiol.2016.48
53. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2014
Also many uncultured eukaryotic groups
5353
54. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Details
• Dowloaded 10,000+ genomes from various
databases (including many I generated)
• 1000+ new genomes
• Searched these genomes for a set
universal homologous genes (ribosomal
proteins) (based on AMPHORA)
• Aligned the sequences of these genes
between species
• Maximum likelihood tree
54
55. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Details
• If you want to do things like this
• Learn biology but also
• Bioinformatics
• Programming
• Data science
• Quantitative biology
• Statistics
55
56. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Reminder
These trees are based on analysis of
ribosomal proteins. They represent only
a small subset of all the genes in a
genome.
56
57. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Microbial Diversity
• We do not have time to cover all of these
groups of microbes in lecture
• These groups barely scratch the surface of
the true diversity
• Examples of Biological Diversity of
Microbes
• Focus on the Big Picture Patterns of This
Diversity
57
59. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Clicker
Which of the following is an example of
universal homology
A: Ether-linked lipids
B: Peptidoglycan
C: Ester-linked lipids
D: Transcription of DNA into RNA
E: Translation of RNA in the nucleus
59
60. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Clicker
Which of the following is an example of
universal homology
A: Ether-linked lipids
B: Peptidoglycan
C: Ester-linked lipids
D: Transcription of DNA into RNA
E: Translation of RNA in the nucleus
60
61. Bacterial Diversity: Gram Positive vs. Negative
61
Outside of cell
Outside of cell
Inside of cell
Inside of cell
Cell
envelope
Cell wall
(peptidoglycan)
Plasma
membrane
Outer membrane
of cell envelope
Periplasmic space
Peptidoglycan layer
Periplasmic space
Plasma
membrane
5 µm
5 µm
Gram Positive
Gram
Negative
62. Bacterial and Archaeal Shapes
Archaea cell membranes
have lipids with fatty acids
linked to glycerol by ether
linkages (a synapomorphy of
archaea):
62
63. Ester Linkages
Bacterial and eukaryotic cell membranes
have lipids with fatty acids connected to
glycerol by ester linkages:
63
64. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Diversity of Form
• Bacteria and archaea way more diverse in
morphology (e.g., size, shape) than many
appreciate
• Morphological diversity in NPAF
eukaryotes also immense (NPAF = non
plant, animal, or fungal)
• Diversity of movement connected to
diversity of form
• Many examples of convergent evolution in
morphology, related features
64
65. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Bacteria: Major Cell Forms
• Among the Bacteria and Archaea, three
shapes are common:
! Sphere or coccus (plural cocci), occur
singly or in plates, blocks, or clusters.
! Rod—bacillus (plural bacilli)
! Helical
• Rods and helical shapes may form chains
or clusters.
65
66. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Bacteria: Other Forms
66
67. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Archaea: Examples of Forms
67
68. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Motility of Vibrio (a member of the Proteobacteria phylum)
68
69. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Corkscrew Movement of Spiraling (A Cyanobacterium)
69
70. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
•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
70
Certium
tenue
Coral symbiont
71. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Chromalveolates: Haptophytes
• Haptophytes
71
Coccolithophores (haptophytes)
can also form immense
blooms in the ocean.
Blooms can reduce the amount
of sunlight that penetrates
deeper waters.
Emiliania huxleyi—one of
smallest unicellular
eukaryotes. May contribute to
global warming through its
metabolism.
72. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Alveolates: Ciliates
72
Movement in a ciliate from the gut of a termite
• All have numerous cilia,
• Most are heterotrophic; very diverse
group.
• Have complex body forms and two
types of nuclei.
73. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Stramenopiles: Diatoms
•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
73
A colony of the diatom,
Bacillaria paradoxa
74. 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.
74
75. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
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.
75
76. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Unikonts: Opisthokonts: Choanoflagellates
•Choanoflagellates are sister to the
animals.
•Some are colonial and resemble a
type of cell found in sponges.
76
The choanoflagellate Salpingoeca sp. feeding
80. 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
80
81. 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
81
Karyo
82. Multicellularity
• Many lineages, not just PAF (plants,
animals and fungi) include multicellular
representatives
• Mechanisms responsible for multicellularity
different in different groups (why might that
be)?
82
83. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Many Fungi Multicellular
83
86. 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
86
Karyo
87. •All are multicellular; some get very large
(e.g., giant kelp).
•The carotenoid fucoxanthin imparts the
brown color.
•Almost exclusively marine.
Stramenopiles: Brown Algae
87
A community of brown algae: The marine kelp forest
88. 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
88
89. Plantae: Red Algae
89
• Most red algae are marine
and multicellular.
• Red pigment is
phycoerythrin.
•Many reproduce with spores
Motile spores from
Purpureofilum
Audouinella pacifica
Spyridia
90. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
The chlorophytes are the
sister group to charophytes
and land plants.
Synapomorphies include
chlorophyll a and b, and
starch as a storage product.
More than 17,000 species;
marine, freshwater, and
terrestrial. Unicellular to large
90
Plantae: Chlorophytes
Movement in the green
alga Volvox
Micrasterias
93. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Photo 26.4 Filaments of photoautotrophic cyanobacteria,
93
94. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
More on Multicellularity
Later in BIS2C
94
95. Diversity of Processes
• Microbes are able to make use of or alter
just about any chemical bond found on
Earth
• This allows a wide range of niches, and a
wide diversity of roles in ecosystems
• Also diverse mechanisms for surviving and
thriving in “harsh” conditions
• Humans and other organisms have taken
advantage of this diversity in many ways
95
96. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
The Unusual
105°C
CH3
CO, 80°CH2S, pH 0, 95°C High salt
CO2 4°Clow pH
96
99. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Functional Diversity
Covered in
Labs 2 and 3 and Lecture 11-13
99
100. Interactions
• Microbes have diverse interactions with
other organisms (both microbes and
macrobes)
• Symbiosis is an intimate association
between at least two different organisms in
which at least one of them benefits
100
102. Alveolates: Apicomplexans
• All parasitic
• Have a mass of organelles at one tip
—the apical complex that help the
parasite enter the host’s cells.
102
Apical complex • Plasmodium falciparum-
Malaria kills 700,000-2,000,000
people per year—75% of them
are African children
103. Slides by Jonathan Eisen for BIS2C at UC Davis Spring 2016
Red tide caused by dinoflagellates (Gonyaulax sp.).
103
104. 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
104
105. 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.
105