Protists

Trumpet-shaped protists (Stentor coeruleus)

Protists
Protist is the informal name of the group of mostly
unicellular eukaryotes

Advances in eukaryotic systematics have caused the

classification of protists to change significantly

Protists constitute a polyphyletic group, and Protista is no

longer valid as a kingdom

Figure 28.1

1 m

Most eukaryotes are single-celled organisms

Protists are eukaryotes
Eukaryotic cells have organelles and are more
complex than prokaryotic cells
The organisms in most eukaryotic lineages are
protists, and
Most protists are unicellular

Endosymbiosis in Eukaryotic Evolution

Protist diversity has its origins in endosymbiosis

Plastid Evolution
Mitochondria arose first through descent from a
bacterium that was engulfed by a cell from an
archaeal lineage

The plastid lineage evolved later from a

photosynthetic cyanobacterium that was engulfed by
a heterotrophic eukaryote

The plastid-bearing lineage of protists evolved into

photosynthetic protists, red and green algae

Figure 28.3

Membranes
are represented
as dark lines
in the cell.
Red alga
Cyanobacterium

Dinoflagellates
Secondary
endosymbiosis

Plastid

1 23
Primary
endosymbiosis

Stramenopiles

Nucleus
Heterotrophic
One of these
eukaryote
membranes
was lost in
red and
green algal
descendants.

Secondary
endosymbiosis

Plastid

Euglenids
Secondary
endosymbiosis

Green alga
Chlorarachniophytes

Figure 28.3a

Primary endosymbiosis
Membranes
are represented
as dark lines
in the cell.
Red alga
Cyanobacterium
1 23

Nucleus
Heterotrophic
One of these
eukaryote
membranes
was lost in
red and
green algal
descendants.
Green alga

Figure 28.3b

Secondary endosymbiosis

Dinoflagellates

Red alga

Plastid

Stramenopiles

Figure 28.3c

Secondary endosymbiosis

Plastid

Euglenids

Green
alga
Chlorarachniophytes

Chlorarachniophytes have a nucleomorph

Inner plastid
membrane

Nucleomorph
Outer plastid
membrane
Nuclear pore-like gap

Figure 28.2a
Parabasalids
Euglenozoans

Excavata

Diplomonads

Stramenopiles

Diatoms

Golden algae
Brown algae

Apicomplexans
Ciliates

SAR clade

Alveolates

Dinoflagellates

Forams
Rhizarians

Cercozoans
Radiolarians

Green
algae

Chlorophytes
Charophytes
Land plants

Archaeplastida

Red algae

Tubulinids

Entamoebas
Nucleariids

Opisthokonts

Fungi
Choanoflagellates
Animals

Unikonta

Amoebozoans

Slime molds

Figure 28.2aa

Parabasalids
Euglenozoans

Excavata

Diplomonads

Stramenopiles

Diatoms
Golden algae
Brown algae

Apicomplexans
Ciliates
Forams

Rhizarians

Cercozoans
Radiolarians

SAR clade

Alveolates

Dinoflagellates

Excavata
5 m

Giardia intestinalis, a diplomonad parasite

The clade Excavata is characterized by its

cytoskeleton

Some members have an excavated feeding groove

This group includes the diplomonads, parabasalids,

and euglenozoans

Diplomonads and Parabasalids

They lack plastids, have modified mitochondria, and
most live in anaerobic environments
Diplomonads
Have reduced mitochondria called mitosomes

Have two equal-sized nuclei and multiple flagella

Are often parasites

Parabasalids
Have reduced mitochondria
Include Trichomonas vaginalis, the pathogen that
causes yeast infections in human females
Flagella

Undulating
membrane

5 m

Euglenozoans
Euglenozoa is a diverse clade that includes
predatory heterotrophs, photosynthetic autotrophs,
mixotrophs, and parasites

The main feature distinguishing them as a clade is a

spiral or crystalline rod inside their flagella

This clade includes the kinetoplastids and euglenids

Long flagellum

Eyespot

Short flagellum

Light
detector

Contractile vacuole
Nucleus
Chloroplast

Euglena (LM)

5 m

Plasma
membrane

Pellicle

Figure 28.8a

Long
flagellum

Eyespot

Contractile
vacuole
Nucleus

Chloroplast

Euglena (LM)

5 m

Plasma
membrane

Video: Euglena

Figure 28.2aa

Parabasalids
Euglenozoans

Excavata

Diplomonads

Stramenopiles

Diatoms
Golden algae
Brown algae

Apicomplexans
Ciliates
Forams

Rhizarians

Cercozoans
Radiolarians

SAR clade

Alveolates

Dinoflagellates

Stramenophila, Alveolata and Rhizaria

SAR Clade

Diatom diversity

50 m

Diverse group of protists defined by DNA

similarities

The SAR clade is a diverse monophyletic

supergroup named for the first letters of its three
major clades stramenopiles, alveolates, and
rhizarians

Excavata
Diatoms
Golden algae
Brown algae

Stramenopiles

Ciliates

Forams
Cercozoans
Radiolarians

SAR clade

Dinoflagellates
Apicomplexans Alveolates

Rhizarians
Archaeplastida
Unikonta

Stramenopiles
The stramenopiles clade includes some of the most
important photosynthetic organisms on Earth

Most have a hairy flagellum paired with a smooth

flagellum

Stramenopiles include diatoms, golden algae, and

brown algae

Stramenophiles

Hairy
flagellum
Smooth
flagellum

5 m

Diatoms
Diatoms are unicellular algae with a unique two-part,
glass-like wall of silicon dioxide

Golden Algae
Golden algae are named for
their color, which results from
their yellow and brown
carotenoids
All golden algae are
photosynthetic, and some are
mixotrophs
Most are unicellular, but
some are colonial

Brown Algae
Brown algae are the largest and most complex
algae

All are multicellular, and most are marine

Brown algae include many species commonly called

seaweeds

Figure 28.12

Blade

Stipe

Holdfast

Alternation of Generations
A variety of life cycles have evolved among the
multicellular algae
The most complex life cycles include an alternation
of generations, the alternation of multicellular
haploid and diploid forms

Heteromorphic generations are structurally

different, while isomorphic generations look similar

The diploid sporophyte produces haploid flagellated

spores called zoospores

The zoospores develop into haploid male and female

gametophytes, which produce gametes

Fertilization of gamates results in a diploid zygote,

which grows into a new sporophyte

Figure 28.13

Haploid (n)
Diploid (2n)
Sporangia

MEIOSIS

10 cm

Sporophyte
(2n)
Developing
sporophyte

Zygote
Mature female
(2n)
gametophyte
(n)
FERTILIZATION

Zoospore

Female

Gametophytes
(n)
Male

Egg

Sperm

Figure 28.13a-1

Haploid (n)
Diploid (2n)

Sporangia

MEIOSIS
Sporophyte
(2n)

Zoospore

Female
Gametophytes
(n)
Male

Egg

Sperm

Figure 28.13a-2

Haploid (n)
Diploid (2n)

Sporangia

MEIOSIS
Sporophyte
(2n)
Developing
sporophyte
Zygote
(2n)

Mature female
gametophyte
(n)
FERTILIZATION

Zoospore

Female
Gametophytes
(n)
Male

Egg

Sperm

Alveolates
Members of the clade Alveolata have membraneenclosed sacs (alveoli) just under the plasma
membrane
The alveolates include
Dinoflagellates

Apicomplexans
Ciliates

Figure 28.14

Alveoli

Alveolate
0.2 m

Flagellum

Dinoflagellates
Dinoflagellates have two flagella and each cell is
reinforced by cellulose plates

They are a diverse group of aquatic phototrophs,

mixotrophs, and heterotrophs

Figure 28.15

Flagella
(a) Dinoflagellate
flagella

3 m

(b) Red tide in the Gulf

of Carpentaria in
northern Australia

Ciliates
They have large macronuclei and small micronuclei
Genetic variation results from conjugation, in which
two individuals exchange haploid micronuclei
Conjugation is a sexual process, and is separate
from reproduction, which generally occurs by binary
fission

Figure 28.17a

Contractile
vacuole

50 m

Cilia

Micronucleus
Macronucleus
(a) Feeding, waste removal, and water balance.

Oral groove
Cell mouth

Food
vacuoles

Paramecium

Rhizarians
Many species in the rhizarian clade are amoebas
Amoebas are protists that move and feed by
pseudopodia, extensions of the cell surface
Rhizarian amoebas differ from amoebas in other
clades by having threadlike pseudopodia
Rhizarians include radiolarians, forams, and
cercozoans

Radiolarians
Marine protists called radiolarians have delicate, symmetrical
internal skeletons that are usually made of silica

Pseudopodia

200 m

Forams
Foraminiferans, or forams, are named for porous, generally
multichambered shells, called tests

Pseudopodia extend through the pores in the test

Many forams have endosymbiotic algae

Figure 28.2a
Parabasalids
Euglenozoans

Excavata

Diplomonads

Stramenopiles

Diatoms

Golden algae
Brown algae

Apicomplexans
Ciliates

SAR clade

Alveolates

Dinoflagellates

Forams
Rhizarians

Cercozoans
Radiolarians

Green
algae

Chlorophytes
Charophytes
Land plants

Archaeplastida

Red algae

Tubulinids

Entamoebas
Nucleariids

Opisthokonts

Fungi
Choanoflagellates
Animals

Unikonta

Amoebozoans

Slime molds

Figure 28.2ab

Green
algae

Chlorophytes

Charophytes
Land plants

Archaeplastida

Red algae

Tubulinids
Entamoebas
Nucleariids

Opisthokonts

Fungi
Choanoflagellates
Animals

Unikonta

Amoebozoans

Slime molds

Red algae and green algae are the closest relatives of

land plants

Plastids arose when a heterotrophic protist acquired

a cyanobacterial endosymbiont
The photosynthetic descendants of this ancient
protist evolved into red algae and green algae

Land plants are descended from the green algae

Red Algae
Accessory pigment called phycoerythrin masks the
green of chlorophyll

Red algae are usually multicellular; the largest are

seaweeds

Red algae are the most abundant large algae in

coastal waters of the tropics

Figure 28.21a

Bonnemaisonia
hamifera

8 mm

Figure 28.21b

20 cm

Dulse (Palmaria palmata)

Nori

Nori

Green Algae
Plants are descended from the green algae

Green algae are a paraphyletic group

The two main groups are the charophytes and the

chlorophytes

Charophytes are most closely related to land plants

Larger size and greater complexity evolved in

chlorophytes by:
1. The formation of colonies from individual cells

2. The formation of true multicellular bodies by cell

division and differentiation (e.g., Ulva)

3. The repeated division of nuclei with no cytoplasmic

division (e.g., Caulerpa)

Figure 28.2a
Parabasalids
Euglenozoans

Excavata

Diplomonads

Stramenopiles

Diatoms

Golden algae
Brown algae

Apicomplexans
Ciliates

SAR clade

Alveolates

Dinoflagellates

Forams
Rhizarians

Cercozoans
Radiolarians

Green
algae

Chlorophytes
Charophytes
Land plants

Archaeplastida

Red algae

Tubulinids

Entamoebas
Nucleariids

Opisthokonts

Fungi
Choanoflagellates
Animals

Unikonta

Amoebozoans

Slime molds

Figure 28.2ab

Green
algae

Chlorophytes

Charophytes
Land plants

Archaeplastida

Red algae

Tubulinids
Entamoebas
Nucleariids

Opisthokonts

Fungi
Choanoflagellates
Animals

Unikonta

Amoebozoans

Slime molds

Unikonts include protists that are closely

related to fungi and animals
The supergroup Unikonta includes animals, fungi,
and some protists
This group includes two clades: the amoebozoans
and the opisthokonts (animals, fungi, and related
protists)

The root of the eukaryotic tree remains controversial

It is unclear whether unikonts separated from other
eukaryotes relatively early or late

Amoebozoans
Amoebozoans are amoeba that have lobe- or tubeshaped, rather than threadlike, pseudopodia
They include slime molds, tubulinids, and
entamoebas

Slime Molds
Slime molds, or mycetozoans, were once thought to be fungi
DNA sequence analyses indicate that the resemblance between
slime molds and fungi is a result of convergent evolution

Slime molds include two lineages, plasmodial slime molds and

cellular slime molds