BIO 12: LEC LE 2

CHAPTER 28: ORIGIN & EVOLUTION OF EUKARYOTES

INTRODUCTION
Even a low-power microscope can reveal a great variety of organisms in a drop of pond
water EUKARYOTES SUPERGROUPS
o Anton von Leeuwenhoek (1632-1723) made the earliest microscope
o Hans Locke made a replica of Leeuwenhoeks microscope in 2007
Protists constitute a polyphyletic group and it is no longer a valid kingdom
Protist informal name of the group of mostly unicellular eukaryotes
o Eukaryotes have organelles and are more complex than prokaryotes
o The organisms in most eukaryotic lineages are protists
o Though most protists are unicellular, there are some colonial and multicellular
species
STRUCTURAL & FUNCTIONAL DIVERSITY
Reproduction some protests reproduce sexually (meiosis, fertilization) while others
reproduce asexually
It is no longer thought that amitochondriates (lacking mitochondria) are the oldest
lineage of eukaryotes
o Many have been shown to have mitochondria and have been reclassified
Classification of organisms has changed significantly due to advances in systematics
o Ultrastructues (ex. SEM)
o Molecular evidence
o Phylogenetic evidence
One hypothesis divides all eukaryotes (including protists) into 4 supergroups

Protists exhibit the most diversity among all other eukaryotes

Single-celled protists can be very complex, as all biological functions are carried out by
organelles in each individual cell
Protists are also the most nutritionally diverse of all eukaryotes
o Photoautotrophs contain chloroplast (ex. Euglena)
o Heterotrophs absorb organic molecules or ingest larger food particles
Ex. Paramecium
Food vacuole can move around, processing food particles
via the digestive enzymes from the lysosomes
Food particle oral groove of cilia pharynx forms a
food vacuole: goes around digesting food change in
acidity
Undigested materials form a temporary break in the
plasma membrane so they can be expelled
Ex. Paramecium bursaria symbiotic relationship with
chlorella, which makes its food
Battle 1: Ciliate (Didinium) engulfs a ciliate (Paramecium) Q: How did protists become so diverse? A: Endosymbiosis in eukaryote evolution
Didinium releases toxicyst to trap o Endosymbiosis
Paramecium panics and releases trichocyst Relationship between two species in which a unicellular
Battle 2: Ciliate engulfs a rotifer organism lives inside the cell/s of another organism (host)
o Mixotrophs photosynthesis + heterotrophic nutrition (ex. Euglena goes Process wherein a unicellular organism engulfs another cell
back and forth to heterotrophic and photoautotrophic nutrition) which becomes an endosymbiont, and then an organelle in the
host cell (cell endosymbiont organelle)
 o Mitochondria and plastids are derived from endosymbiosis
Mitochondria evolved first by endosymbiosis of an alpha
proteobacterium by a cell from an archaeal lineage
Plastids evolved later by endosymbiosis of a photosynthetic
cyanobacterium by a heterotrophic eukaryote; evolved further
into red/green photosynthetic algae
o The ancestral host cell may have been an archaean or protoeukaryote,
from a lineage related to but diverged from archaeal ancestors
o Secondary endosymbiosis ingested in the food vacuole of heterotrophic
eukaryotes and became endosymbionts themselves (ex. red/green algae)
Red algae dinoflagellates, stramenopiles
Green algae chlorarachniophytes, euglenids
Nucelomorph vestigial nucleus of the engulfed cell
o Evidences of cyanobacterial origins of plastids in protists (red/green
algae)
Have 2 membranes
Similar transport proteins in the membranes
I. EXCAVATA
Characterized by its cytoskeleton; some have a feeding groove
Diplomonads and parabasalids were: (a) once thought to be amitochondriates; now
have modified mitochondria, (b) lack plastids, (c) live in anaerobic environments;
generates energy anaerobically
1. Diplomonads
o Mitosomes reduced mitochondria
o Have two equal-sized nuclei and multiple flagella
Prokaryotic flagella differs from eukaryotic flagella in that they are
filaments composed of globular proteins attached to the cell surface
o Often parasites
Ex. Giardia intestinalis causes giardiasis, an infection caused by
trophozoites in the small intestine
2. Parabasalids
o Hydrogenosomes reduced mitochondria that generate some energy
anaerobically
o Ex. Trichomonas vaginalis pathogen that causes yeast infections in
females and trichomoniasis STD
Trichomonas infections: pelvic inflammatory disease, adverse
pregnancy outcomes, infertility, prostate cancers, increased
HIV-1 transmission
o Ex 2. Trichonympha symbiotic parabasalid living in gut of termites to
enable then to digest cellulose in wood
3. Euglenozoans
o Diverse predatory/photosynthetic heterotrophs, mixotrophs, parasites
o Have a spiral/crystalline rod inside each flagella
o Flagella and cilia have a common structure
Microtubules covered by plasma membrane
Basal body for anchorage
Dynein motor protein, which drives the bending movements
a. Kinetoplastids
Kinetoplast organized mass of DNA inside a single mitochondrion;
organelle within an organelle
Structure
Axoneme (upper) for locomotion; has 9+2 configuration
Kinetosome (lower) for anchorage; has 9 peripheral
triplets and no central microtubule
Consumers free-living species are consumers of prokaryotes in
freshwater (ex. Bodo saltans - bacteriovore), marine, and moist
terrestrial ecosystems
Parasites
Ex 1. Trypanosoma brucei gambiense/rhodesiense/brucei
causes sleeping sickness in humans
o Note: only the first two subspecies cause death
o Occurs via bite of tsetse fly (Glossina spp.) which
travels via the CSF to the brain
o Symptoms: falling asleep anytime and anywhere,
mental dumbness
o If untreated, leads to comatose
o Medecins Sans Frontieres (MSF) has a sleeping
sickness ward in Sudan
Ex 2. Trypanosoma cruzi causes Chagas disease by blood
sucking insects
Trypanosomes bait-and-switch defense to evade immune
responses
Trypanosomes aka hemoflagellates surround RBCs
A trypanosome produces millions of copies of a single
protein
They evade the hosts immune responses by switching
surface proteins
Before the immune system can recognize the protein and
mount the attack, new generation of trypanosomes makes
 millions of copies of a different surface protein
These frequent changes prevent the host from developing
immunity
b. Euglenids
Have one or two flagella that emerge from a pocket at one end of
the cell
Some species can be mixotrophic
Ex. Euglena commonly found in pond water
II. SAR CLADE
b.
Diverse
Named for the first letters of the 3 major clades: (S)tramenopiles, (A)lveolates,
(R)hizarians
One of the most controversial supergroups
1. Stramenopiles
2. Alveolates
o Alveoli membrane-enclosed sacs just under the plasma membrane
a. Dinoflagellates (flagellates)
Structure 2 flagella, each cell reinforced by cellulose plates
Aquatic phototrophs, mixotrophs, heterotrophs
Abundant components of marine/freshwater phytoplankton
Reason why coral reefs are biologically diverse and productive
Toxic red tides are caused by dinoflagellate blooms
Ex 1. Pyrodinium bahamense var. compressum causes
HABs in the Philippines; first discovered in Red Sea (1983)
and got to the Philippines via the opening of the Suez
Canal, which was developed by Ferdinand de Lesseps and
links the Mediterranean Sea to Red Sea
Ex 2. Pfiesteria shumwayae causes toxic red tides
Ex 3. Pfiesteria piscicida causes fish kills in the North
Atlantic Ocean; life cycle has 24 stages; (a) flagellated
stage suck out juices, (b) amoeboid stage feed on dead fish
Ex 4. Ceratium
Ex 5. Gonyaulax
Ex 6. Noctiluca scintillans bioluminescent dinoflagellates,
non-photosynthetic, non-motile, microbivores
Ex 7. Orbitolites vertebralis shows algal windows
Ex 8. Symbiodinium
Planktonic
Mastigote swimming form of symbiont turns into
coccoid cells in the gastrodermis move to the inside of
the gastrovascular cavity of the host
Zooxanthellae become endosymbionts; share organic
compounds with their host cell
Gastrodermis of Cnidarians (host) provides: (a) protection
from herbivores while they are a mastigote, (b) raw
materials for photosynthesis (CO2, NH4, PO4)
Symbiodinium (parasite) in return provides: fatty acids,
lipid droplets, gas, glucose, glycerol
Apicomplexans (parasites)
Apicomplexan named after the apex end of the sporozoite
cointains complex of organelles for penetrating host cells/tissues
Apicoplastid from red alga; absent in gregarines, Colpodellids,
Cryptosporidium spp.
Parasites of animals, some cause serious human diseases
Not photosynthetic
No means of locomotion
Most have sexual and asexual stages that require two or more
different host species for completion
Ex 1. Chromera velia free-living, coral associated, has
photosynthetic apicoplast
Ex 2. Plasmodium causes malaria
Species present in the Philippines, from most to least
virulent: Plasmodium falciparum > Plasmodium malariae >
Plasmodium ovale > Plasmodium vivax
Requires both female mosquito and human to complete its
life cycle
Anopheles gambiae top vector of Plasmodium
68 transmit malaria / 460 or 422 Anopheles / 3000-3500
species of mosquitoes
Has three invasive stages:
i. Ookinete (in mosquito) without apical complex
ii. Sporozoite (in mosquito) from sporogony; infectious
cells that they go as, to spread throughout their host
iii. Merozoite (in human) with apical complex
Approximately 900,000 people die each year from malaria
Life cycle of Plasmodium falciparum
In human:
 1. Female transfers sporozoite from saliva
2. Sporozoites break through the skin to the blood
stream and blood vessels
3. Sporozoites enter the liver cells
4. Shizogony sporozoites multiply in the liver cells via
multiple fission
5. Merozoites escape from the liver cells
6. Merozoites penetrate RBCs produce merozoin toxin
and digest hemoglobin
7. RBCs burst and release the merozoin toxin
8. Merozoites become gametocytes, which give rise to
gametes
In mosquitoes:
9. Female mosquito sucks RBCs with gametocytes so
they go to the intestine
Microgamete = sperm equivalent
Macrogamete = egg equivalent
10. Microgamete fertilizes macrogamete to produce a
zygote
11. Zygote goes to the stomach and becomes an ookinete
(2n)
12. Ookinete passes through the peritrophic membrane
(chitinous tough lining from stomach to hemocoelom)
13. Ookinete becomes an oocyst
14. Sporogony happens in the oocyst to produce many
sporozoites
15. Sporozoites break out from oocyst
16. Sporozoite goes to salivary glands and waits there
until the female bites someone to transfer those
sporozoites to
Difficulties in controlling malaria
Different species of Plasmodium
Different stages in the life cycle
Intracellular stays within our own cells, which our
immune system cannot attack
Has various stains different surface antigens
Alters surface chemistry of infected blood cells RBCs
stick to the lining of the blood vessels and cannot be
brought to the spleen for destruction, will stay as PfEMP1
instead
Vector mosquitoes have persistence of breeding places
(fresh waters)
Vector mosquitoes have different susceptibilities to
insecticides; resistant to DDT
Human host are exposed to mosquito bites; even in tiny
places, mosquitoes find ways
Recent approaches
Effective genes inserted into the mosquitos genome to
prevent penetration of ookinete to peritrophic membrane:
SM1, PLA2
Single-chain antibody can stop sporozoites from going to
salivary glands
Genetically modified (?)
Celebrities who help raise malaria awareness
Bill Gates
Stephen Curry
c. Ciliates (animal-like protists)
Large, varied group of protists that are mostly predators
Cilia named for their use of cilia to move and feed
Cilia usually cover the whole structure, but some have
reduced or modified cilia (ex. Stylonychia, Oxytricha
trifallax, Vorticella)
Cirri bundles of cilia
Membranelle fine sheaths of cilia
Food vacuole engulfs a food item via phagocytosis
Contractile vacuole
Uses: osmoregulation, volume regulation
Only present in freshwater protists (ex. Paramecium,
Euglena); totally absent in marine and parasitic protists
Dimorphic have large macronuclei and small micronuclei
Macronuclei for conjugation
Micronuclei for vegetative functions and for regulation
of ciliate functions
Monomorphic some protists may have only one kind of nucleus
Ex. Balantidium coli parasitic ciliate
Found in stool; causes balantidiasis (diarrhea)
Only known ciliate affecting humans; largest protist
parasite in humans; prevalent among Filipinos
You can get this if you stay with pigs
Conjugation
Exclusive to ciliates
Two conjugants (must be of same species) exchange
haploid micronuclei
Conjugation is a sexual process separate from
 reproduction (via transverse binary fission)
i. Conjugants 2 compatible mates (same species,
same syngens, different mating types) come together
ii. Conjugation bridge temporary cytoplasmic union
iii. Meiosis of micronuclei
iv. Exchange of micronuclei
v. Ex-conjugants conjugants separate
vi. One of the two ex-conjugants undergoes
micronuclear fusion original macronucleus
disintegrates new macronuclei formed from
micronuclei divide by binary fission
Product 4 individuals all genetically different
(conjugation leads to genetic variation)
Why do ciliates conjugate? (a) Conjugation rejuvenates
aging ciliates, (b) Conjugation is necessary during
unfavorable conditions like starvation
Ex 1. Paramecium caudatum alternate between
conjugation and asexual reproduction via transverse
binary fission
Ex 2. Nassula ornate freshwater ciliate exhibiting
conjugation
3. Rhizarians
o DNA evidence supports Rhizaria as a monophyletic clade
o Mostly amoebas but also includes radiolarians, forams, cercozoans
o Pseudopodia cellular extensions that allow amoebas to move and feed
Rhizarian amoebas differ from others by having thread-like
pseudopodia (axopodia)
a. Radiolarians
Have delicate, symmetrical internal skeletons made of silica
Pseudopodia - to engulf microorganisms via phagocytosis; radiate
from central body
b. Foraminiferans/Forams
Foramina pores in multichambered shell made of calcium
carbonate (test); where pseudopodia extent
Pseudopodia extend through pores in the test
Many forams have endosymbiotic algae (ex. Peneroplis pertusus has
an endosymbiotic alga, Porphyridium purpureum)
Examples: (1) Baculogypsina, (2) Globigerina, (3) Cribrinonion lene,
Foram tests in marine sediments form an extensive fossil record
Some fossil species indicate oil deposits
Researchers can use measures of magnesium content in
fossilized forams to estimate changes in ocean
temperature over time
c. Cercozoans
Include most amoeboid and flagellated protists with filopodia
Common in marine, freshwater, and soil ecosystems
Mostly heterotrophs (parasites, predators)
Ex. Paulinella chromatophora
Autotroph with chromatophore for photosynthesis
Chromatophores evolved from a different cyanobacterium
than the plastids of other photosynthetic eukaryotes
Prochlorococcus cyanobacteria that might have been the
endosymbiont
Has a single pseudopodium for locomotion
III. ARCHAEPLASTIDA
Red and green algae are the closest relatives of land plants
Land plants are descended from the green algae
IV. UNIKONTA
Root of the embryonic tree remains controversial; it is unclear whether unikonts
separated from other eukaryotes relatively early or late
1. Amoebozoans
o Amoebas that have lobe/tube-shaped pseudopodia (lobopodia) rather
than thread-like
o Ex. Chaos carolinese has green alga prey, Pandorina morum
a. Slime molds (not protozoans)
b. Tubulinids
Commonly unicellular protists in soil, freshwater, or marine
environments
Mostly heterotrophic and actively seek to consume bacteria and
other protists
Ex 1. Arcella testate amoeba
Ex 2. Naegleria fowleri soil amoeba; when in contact with warm
 water, may enter the nose of the swimmer travels to nasal o Pseudopfiesteria shumwayae
passage olfactory nerves skull brain causes primary o Pfiesteria piscicida
amoebic meningoencephalitis (PAM) o Phytophora ramorum causes sudden oak death
c. Entamoebas o Phytophora infestans causes potato late blight, which contributed
Parasites of vertebrates and some invertebrates to the Irish famine in the 19th century
Ex 1. Entamoeba histolytica causes amoebic dystentery, 2. Photosynthetic protists
the third leading cause of human death due to eukaryotic Many are primary producers that obtain energy from the sun
parasite In aquatic environments
Ex 2. Entamoeba gingivalis commensal in the mouth o Photosynthetic protists and prokaryotes are the main primary
2. Opisthokonts producers
a. Animals o Photosynthetic protists are limited by nutrients
b. Fungi These populations can explode (as HABs) when limiting nutrients are in excess
c. Choanoflagellates Biomass of photosynthetic protists has declines as sea surface temperature
Held together by ECM and intracellular bridges has increased
Intermediate between animals and fungi Upwelling delivers nutrients from the ocean bottom
o Growth of phytoplankton communities relies on the nutrients
REPRODUCTION OF PROTISTS acquired from this
o Warm surface water acts as a barrier to upwelling
ASEXUAL
If sea surface water continues to warm due to global warming, this could have
A. Binary Fission
large effects on
a. Transverse fission in ciliates
o Marine ecosystems
b. Longitudinal fission in euglenids, kinetoplastids, diplomonads
o Fishery yields
c. Oblique fission in dinoflagellates
o The global carbon cycle
B. Multiple Fission
a. Schizogony in sporozoites and merozoites of Plasmodium (human host)
b. Sporogony in oocyst of Plasmodium (mosquito host)

SEXUAL
A. Conjugation in ciliates only; same species and variety (syngen) but different
mating types (ex. Spirostomum)

ECOLOGICAL ROLES OF PROTISTS

Protists are found in diverse aquatic and moist terrestrial environments

Protists play two key roles in their habitats: symbionts and producers

1. Symbiont protists
Benefit their host
o Dinoflagellates nourish the coral polyps that build reefs (ex.
zooxanthellae in the gastrodermis of corals)
o Wood-digesting protists inhabit the gut of termites (ex. parabasalid)
Parasitic or predatory
o Plasmodium causes malaria