2019/1/14

Protist Parasites Most eukaryotes are single‐celled 
organisms
• Protists are eukaryotes
• Eukaryotic cells have organelles and are more complex 
than prokaryotic cells
• Most protists are unicellular, but there are some 
colonial and multicellular species

Three domains of Life

Campbell Biology, 9e. 2011

Structural and Functional Diversity in 
Protists
• Protists exhibit more structural and functional diversity 
than any other group of eukaryotes

• Some protists reproduce asexually, while others 
reproduce sexually, or by the sexual processes of meiosis 
and fertilization
多細胞生物
(Metazoa) • Single‐celled protists can be very complex, as all 
biological functions are carried out by organelles in each 
individual cell…
Most eukaryotes are single‐celled organisms!

1
 2019/1/14

• Protists, the most nutritionally diverse of all eukaryotes,  • Classification of Protists:

include
– Photoautotrophs, which contain chloroplasts – Complexity and diversity of protists makes
– Heterotrophs, which absorb organic molecules or  them difficult to classify
ingest larger food particles
• Many classification schemes proposed
Parasitic!
– Mixotrophs, which combine photosynthesis and 
heterotrophic nutrition
• None has broad support
• Cannot be classified as plants, animals, or fungi
• Not monophyletic
– Do not all belong to the same evolutionary lineage
• Classified into six supergroups
– Taxonomic group below domain and above kingdom
Euglena Amoeba Plasmodium

紅藻、綠藻、植物

頂複器蟲、雙鞭毛藻、
纖毛蟲

褐藻、金黃藻、矽藻

雙滴蟲、毛滴蟲
眼蟲、錐形蟲

變形蟲

真菌、動物

放射蟲、有孔蟲

2
 2019/1/14

Evolutionary Relationships Between the Eukaryotic  Supergroups of Parasitic Protist

Supergroups

Ch10
Ch8, 9 Ch10
Ch7

Ch6 Ch5

Endosymbiosis theory
Endosymbiosis in Eukaryotic Evolution Nucleus

• There is now considerable evidence that much protist  Engulfing of oxygen-using

Ancestor of
prokaryote, i.e. Proteobacteria
diversity has its origins in endosymbiosis eukaryotic cells
(host cell)

Mitochondrion
• Endosymbiosis is the process in which a unicellular 
organism engulfs another cell, which becomes an  Engulfing of photosynthetic
endosymbiont and then organelle in the host cell  prokaryote, i.e. Cyanobacteria

– Mitochondria evolved by endosymbiosis of an aerobic 
prokaryote Chloroplast

– Plastids evolved by endosymbiosis of a photosynthetic  Non-photosynthetic eukaryote

cyanobacterium

Photosynthetic eukaryote

Figure 28.2a

Membranes 紅藻、綠藻、植物
are represented
as dark lines in
Red alga
the cell.
Cyanobacterium 頂複器蟲、雙鞭毛藻、
1 2 纖毛蟲

Primary 褐藻、金黃藻、矽藻
endosymbiosis

雙滴蟲、毛滴蟲
眼蟲、錐形蟲
Heterotrophic
eukaryote
變形蟲

真菌、動物

Green alga 放射蟲、有孔蟲

3
 2019/1/14

Figure 28.2c Figure 28.2b

Plastid

Dinoflagellates
雙鞭毛蟲

Secondary Plastid
endosymbiosis
Secondary
endosymbiosis
Apicomplexans
Euglenids 頂複器蟲
眼蟲
Secondary Red alga
endosymbiosis

Green alga

Chlorarachniophytes
絲足蟲藻 Stramenopiles
不等鞭毛類

Apicomplexans (頂複器蟲) Life Cycle of Plasmodium vivax

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

• Non‐motile obligate parasites female
gamete
Sexual phase
in mosquito

• Form spores
male
gamete
food
canal

• Most serious parasitic disease of humans is malaria
– Plasmodium spp. (瘧原蟲) zygote

sporozoite
1. In the gut of a female Anopheles

• Infects 350‐500 million people each year, of whom more than a  mosquito, gametes fuse, and the

zygote undergoes many divisions
to produce sporozoites, which salivary

million die of the infection migrate to her salivary gland. glands

• Transmitted by mosquito 2. When the mosquito bites a human,

the sporozoites pass from the mosquito
salivary glands into the bloodstream
and then the liver of the host.

– Toxoplasma (弓漿蟲)
3. Asexual spores

• Causes toxoplasmosis
6. Some merozoites
(merozoites) produced become gametocytes,
in liver cells which enter the blood-
enter the blood- liver stream. If taken up
stream and then the cell

• Infects cats and humans
by a mosquito, they
red blood cells, where become gametes.
they feed as trophozoites. gametocytes

• Causes birth defects and mental retardation when pregnant  Asexual phase

in humans

women are exposed 4. When the red blood

cells rupture, merozoites
invade and reproduce
5. Merozoites and
toxins pour
into the blood-

– Cryptosporidium (隱孢子蟲)
asexually inside stream when
new red blood the red blood
cells. cells rupture.

22

Life Cycle of Toxoplasma gondii

Apicomplex

An unique organelle
with 4 layers of
membrane, vital to
parasite’s survival… Toxoplasma

4
 2019/1/14

Apicoplast TEM images of Sarcocystis 2nd Endosymbiosis

Tomova et al. 2006

Apicomplexa
Dinoflagellata
Ciliates…
Mitochondrion

Cyanobacteria

Why 4 layers?
Where did it come from?
Facchinelli and Weber 2011

Type II fatty acid

biosynthesis

Nature. 1997
herbicides

Animal or Plant- Apicomplexan

plant plastid plastid

Enzymes encoded from Plastid‐derived genome 
Waller et al. 2003
Goodman & McFadden 2007

Cryptosporidium parvum causes Cryptosporidiosis

隱孢子蟲 Cryptosporidium parvum 隱孢子
蟲
Scientific classification
Kingdom: Chromalveolata

Phylum: Apicomplexa

Class: Conoidasida
Sporozoite
Subclass: Coccidiasina apical complex
of C. parvum
Order: Eucoccidiorida

Family: Cryptosporidiidae
No apicoplast !!!
Genus: Cryptosporidium nucleus
Species: C. parvum mitochondrion

5
 2019/1/14

Science 2004

Zhu et al. 2002

Obornik et al., 2012,

Secondary endosymbiosis

ENDOSYMBIOTIC GENE TRANSFER (EGT)

Gain some,
lose some.

Chan et al. 2010

6
 2019/1/14

Cryptosporidiosis surveillance 2005‐008, USA Cryptosporidium parvum 隱孢子
蟲

No apicoplast, no target?

Genes associate with Metabolism Pathway Select 14 enzymes involving 
glycolysis in Crypto
Hexokinase (HK)
Phosphoglucomutase (PGluM1, 
PGluM2)
Phosphoglucose isomerase (PGI)
PPi‐dependent phosphfructokinase
(PPi‐PFK1 & PPi‐PFK2) 
Aldolase
Triosephosphate isomerase (TIM)
Glycerol 3‐phosphate 
dehydrogenase (GPDH)
Glyceraldehyde 3‐phosphate 
dehydrogenase (GAPDH)
Phosphoglycerate kinase (PGK) 
Phosphoglycerate mutase (PGM)
Enolase
Pyruvate kinase (PK)
Phosphoenolpyruvate carboxylase 
(PEPCL)
Malic enzyme (ME)

Phosphoglycerate kinase (PGK)* Similar to organismal tree (sort of.)

■ Bactria
■ Archea
■ Other Protist
■ Apicomplexans
■ Animals
■ Plants
● Clade with support >80%
based on MP, ML, or BI.

7
 2019/1/14

Mitochondria associated Unresolved Plastid-associated Glycolysis Enzymes

Plastid‐associated Glycolysis Enzymes 
Taxonomic  Groups Note
Enzyme Api‐ Protist Fungi Plant Animal Archae Bacteria
Potential drug targets~ 
complexan
Hexokinase (HK) Plasmo Toxo Protists Plant  Animal No  No  Only in eukaryotes
(Trypano‐ homolog homolog Enzyme Potential  Enzyme Potential 
some)
Phosphoglucomutase Toxo Ciliates (Plasmo) 2 types of PGluM, Plasmo has 
Target Target
(PGluM1, PGluM2) typeII; Eukaryote GluM may 
come from cyanobacteria  HK GAPDH
(Choroplas origin)
*Phosphoglucose Plasmo Toxo Plant alpha‐ Possible EGT; Gp and other  
isomerase (PGI) (cytosolic) proteo and  apicomplexa have plant  PGluM1,
PGK
other  (cytosolic) type, likely most close  PGluM2
bacteria to alpha‐proteobacteria
*PPi‐dependent  Plasmo Protist Unique bacteria type,  Possible 
phosphfructokinase No Toxo No Giardia LGT, belong to GroupII, clade  PGI PGM
(PPi‐PFK1 & PPi‐PFK2)  No Entamo “Long”  group; see Muller et al., 
2001 PPi-PFK1,
*Aldolase Plasmo Toxo Other  Plant  Proteo‐ Giardia and Entamoeba have  Enolase
Protists (cytosolic)  bacteria +  different type Aldolase;  PPi-PFK2
plant  Kinetoplastid has proteo‐
plastid type bacteria type; Cp and  Aldolase PK
Apicomplexa Aldo may come 
from mitochondria 
Triosephosphate
isomerase (TIM)
Plasmo
No Toxo
Other 
Protists
Pretty much follow organismal 
tree
TIM PEPCL

Glycerol 3‐phosphate  Plasmo, Plant  Animals (kidneto‐ Apicomplexans most close to 

dehydrogenase (GPDH) No Toxo plastid) plant, then animal and archea; 
GPDH ME
similar to organismal tree
Glyceraldehyde 3‐ Plasmo, Toxo Other  Need to include plastid type of 
phosphate  protists Taxo and other protists; redo 
dehydrogenase  with  analyses
(GAPDH) cytosolic 
type 

紅藻、綠藻、植物

頂複器蟲、雙鞭毛藻、
纖毛蟲

褐藻、金黃藻、矽藻

雙滴蟲、毛滴蟲
眼蟲、錐形蟲

變形蟲
Amitochondrial Protists

真菌、動物

放射蟲、有孔蟲

8
 2019/1/14

Hedges 2002

Martin 2010

H:  Hydrogenosome
M: mitosomes
Summary
• The apicoplast of apicomplxan were gained through the secondary
endosymbiosis.
• The missing of apicoplast in gregarines and cryptosporidium should
be secondary due to their adaptation to parasitism.
• Enzymes participate glycolysis in Cryptosporidium parvum
apparently have different phylogenetic affinities.
• The plastid-derived enzymes of animal protist parasites
(Plasmodium, Toxoplasma, Cryptosporidium, Coccidia…) present
potential drug targets for therapies.
• Genes involving metabolic functions may experience frequent
horizontal gene transfer (HGT) or endo-symbiotic gene transfer
(EGT) in early eukaryotic evolution.

Martin 2010

9
 2019/1/14

Not just what you inherited… 

凡走過必留下痕跡
At least for the evolution of unicellular organisms.

END

10