28

KEY CONCEPTS
Protists

28.1 Most eukaryotes are single-celled

organisms p. 594

28.2 Excavates include protists with

modified mitochondria and protists
with unique flagella p. 597

28.3 SAR is a highly diverse group

of protists defined by DNA
similarities p. 601

28.4 Red algae and green algae are the

closest relatives of plants p. 609

28.5 Unikonts include protists that

are closely related to fungi and
animals p. 611
Figure 28.1 Knowing that prokaryotes are extremely small, you might assume that
28.6 Protists play key roles in ecological this image shows six prokaryotes and one much larger eukaryote. But in fact, the only
communities p. 614 prokaryote is the organism immediately above the scale bar—the rest are members
of diverse, mostly unicellular groups of eukaryotes informally known as protists.
Study Tip
Label a drawing: Protist life cycles vary
greatly. To make sense of this variation, What gave rise to the great diversity of protists,
it can be helpful to identify when mitosis and how have their lineages diverged over time?
occurs in different life cycles, as shown
here for a portion of Figure 28.14. Identify Much of protistan diversity can be traced to endosymbiosis:
the stages at which mitosis occurs in
Figures 28.14, 28.18, 28.25, and 28.28. Ancestral
cell Engulfed aerobic
bacterium
Early eukaryote
(a heterotroph)
MEIOSIS
Mitochondrion
Zoospore
Mitosis

FERTILIZATION

Female Mitosis
gametophyte Over time, early heterotrophic and
photosynthetic eukaryotes have given
Engulfed rise to four highly diverse supergroups
photosynthetic of eukaryotes:
bacterium
Go to Mastering Biology Excavata

For Students (in eText and Study Area) Early photosynthetic

• Get Ready for Chapter 28 eukaryote SAR
• Animation: Life Cycle of a Malaria Parasite Plastid
For Instructors to Assign (in Item Library) Archaeplastida
• Scientific Skills Exercise: Interpreting
Comparisons of Genetic Sequences
Unikonta
• Misconception Questions 136–140

593
 CONCEPT 28.1 For example, most protists are unicellular, although there
are some colonial and multicellular species. Single-celled
Most eukaryotes are single- protists are justifiably considered the simplest eukaryotes,
but at the cellular level, many protists are very complex—
celled organisms the most elaborate of all cells. In multicellular organisms,
Protists—along with plants, animals, and fungi—are eukary- essential biological functions are carried out by organs.
otes; they are in domain Eukarya, one of the three domains Unicellular protists carry out the same essential functions,
of life. Genetic and morphological studies have shown that but they do so using subcellular organelles, not multicellu-
some protists are more closely related to plants, fungi, or ani- lar organs. The organelles that protists use are mostly those
mals than they are to other protists. As a result, the kingdom discussed in Figure 6.8, including the nucleus, endoplasmic
in which all protists once were classified, Protista, has been reticulum, Golgi apparatus, and lysosomes. Certain protists
abandoned, and various protist lineages are now recognized also rely on organelles not found in most other eukaryotic
as major groups in their own right. Most biologists still use cells, such as contractile vacuoles that pump excess water
the term protist, but only as a convenient way to refer to from the protistan cell (see Figure 7.13). One group of dino-
eukaryotes that are not plants, animals, or fungi. flagellate protists has the most complex subcellular structure
Unlike the cells of prokaryotes, the cells of protists and known: a unique, eye-like organelle called an ocelloid with
other eukaryotes have a nucleus and other membrane- components resembling the lens and retina in the eyes of
enclosed organelles, such as mitochondria and the Golgi vertebrates and some other animals (Figure 28.2).
apparatus. Such organelles provide specific locations where Protists are also very diverse in their nutrition. Some
particular cellular functions are accomplished, making the protists are photoautotrophs and contain chloroplasts. Some
structure and organization of eukaryotic cells more complex are heterotrophs, absorbing organic molecules or ingesting
than those of prokaryotic cells. Eukaryotic cells also have a larger food particles. Still other protists, called mixotrophs,
well-developed cytoskeleton that extends throughout the combine photosynthesis and heterotrophic nutrition.
cell (see Figure 6.20). The cytoskeleton provides the struc- Photoautotrophy, heterotrophy, and mixotrophy have all
tural support that enables eukaryotic cells to have asym- arisen independently in many different protist lineages.
metric (irregular) forms, as well as to change in shape as Reproduction and life cycles also are highly varied among
they feed, move, or grow. In contrast, prokaryotic cells lack protists. Some protists are only known to reproduce asexu-
a well-developed cytoskeleton, thus limiting the extent to ally; others can also reproduce sexually or at least employ the
which they can maintain asymmetric forms or change shape sexual processes of meiosis and fertilization. All three basic
over time. types of sexual life cycles (see Figure 13.6) are represented
We’ll survey the diversity of eukaryotes throughout the among protists, along with some variations that do not quite
rest of this unit, beginning in this chapter with the protists. fit any of these types. We will examine the life cycles of sev-
As you explore this material, bear in mind that eral protist groups later in this chapter.
• the organisms in most eukaryotic lineages are protists, and
• most protists are unicellular. Endosymbiosis in Eukaryotic Evolution
Thus, life differs greatly from how most of us commonly What gave rise to the enormous diversity of protists that exist
think of it. The large, multicellular organisms that we know today? There is abundant evidence that much of protistan
best (plants, animals, and fungi) are the tips of just a few diversity has its origins in endosymbiosis, a relationship
branches on the great tree of life (see Figure 26.21).
c Figure 28.2 The
Mastering Biology HHMI Video: Seeing the eye-like ocelloid of “Retina”
Invisible: Van Leeuwenhoek’s First Glimpses of Erythropsidinium, a
the Microbial World dinoflagellate. This “Lens”
complex, subcellular
structure resembles the
Structural and Functional Diversity camera-like eye found
in Protists in vertebrates and
some molluscs. In some
Given that they are classified in a number of different dinoflagellates, the ocelloid
groups, it isn’t surprising that few general characteristics may function in prey
of protists can be cited without exceptions. In fact, pro- detection.
tists exhibit more structural and functional diversity than
the eukaryotes with which we are most familiar—plants,
animals, and fungi.

594 UNIT FIVE The Evolutionary History of Biological Diversity

between two species in which one organism lives inside the mitochondrial genes (mtDNA) to those found in major clades
cell or cells of another organism (the host). In particular, as of bacteria. In the Scientific Skills Exercise, you will interpret
we discussed in Concept 25.3, structural, biochemical, and one such set of DNA sequence comparisons. Collectively,
DNA sequence data indicate that mitochondria and plastids such studies indicate that mitochondria arose from an alpha
are derived from bacteria that were engulfed by the ancestors proteobacterium (see Figure 27.17). Results from mtDNA
of early eukaryotic cells. The evidence also suggests that mito- sequence analyses also indicate that the mitochondria of
chondria evolved before plastids. Thus, a defining moment in protists, animals, fungi, and plants descended from a single
the origin of eukaryotes occurred when a host cell engulfed a common ancestor, thus suggesting that mitochondria arose
bacterium that would later become an organelle found in all only once over the course of evolution. Similar analyses
eukaryotes—the mitochondrion. provide evidence that plastids descended from a single com-
To determine which bacterial lineage gave rise to mito- mon ancestor—a cyanobacterium that was engulfed by a
chondria, researchers have compared the DNA sequences of eukaryotic host cell.

Scientific Skills Exercise

Interpreting Comparisons of Genetic

Sequences
Which Bacteria Are Most Closely Related to
Mitochondria? Early eukaryotes acquired mitochondria by
endosymbiosis: A host cell engulfed an aerobic bacterium that c Wheat, used
persisted within the cytoplasm to the mutual benefit of both as the source of
cells. In studying which living bacteria might be most closely mitochondrial RNA
related to mitochondria, researchers compared DNA sequences
encoding ribosomal RNA (rRNA), a component of ribosomes. summarizes the comparison of 617 nucleotide positions from the
Ribosomes perform critical cell functions. Hence, genes encod- gene sequences. Each value in the table is the percentage of the
ing rRNA are under strong selection and change slowly over 617 nucleotide positions for which the pair of organisms have
time, making them suitable for comparing even distantly related the same base. Any positions that were identical across all six
species. In this exercise, you will interpret some of the research RNA-encoding gene sequences were omitted from this compari-
data to draw conclusions about the phylogeny of mitochondria. son matrix.
How the Research Was Done Researchers isolated and cloned
DNA sequences from the gene that codes for the small-subunit INTERPRET THE DATA
rRNA molecule for the wheat (a eukaryote) mitochondrion and 1. First, make sure you understand how to read the comparison
for five bacterial species: matrix. (a) Find the cell that represents the comparison of
• Wheat, used as the source of genes encoding mitochondrial rRNA C. testosteroni and E. coli. What value is given in this cell? (b) What
• Agrobacterium tumefaciens, an alpha proteobacterium that does that value signify about the comparable gene sequences in
lives within plant tissue and produces tumors in the host those two organisms? (c) Explain why some cells have a dash rather
• Comamonas testosteroni, a beta proteobacterium than a value. (d) Why are some cells shaded gray, with no value?
• Escherichia coli, a well-studied gamma proteobacterium that 2. Why did the researchers choose one plant mitochondrion and
inhabits human intestines five bacterial species to include in the comparison matrix?
• Mycoplasma capricolum, a gram-positive mycoplasma, which is
the only group of bacteria lacking cell walls 3. (a) Which bacterium has an rRNA-encoding gene that is most
• Anacystis nidulans, a cyanobacterium similar to that of the wheat mitochondrion? (b) What is the
significance of this similarity?
Data from the Research Gene sequences encoding rRNA for
the wheat mitochondrion and the five bacteria were aligned Instructors: A version of this Scientific Skills Exercise can be
and compared. The data table below, called a comparison matrix, assigned in Mastering Biology.

Wheat mitochondrion A. tumefaciens C. testosteroni E. coli M. capricolum A. nidulans

Wheat mitochondrion – 48 38 35 34 34
A. tumefaciens – 55 57 52 53
C. testosteroni – 61 52 52
E. coli – 48 52
M. capricolum – 50
A. nidulans –
Data from D. Yang et al., Mitochondrial origins, Proceedings of the National Academy of Sciences USA 82:4443–4447 (1985).

CHAPTER 28 Protists 595

 Progress has also been made toward identifying the host endosymbiont—a photosynthetic cyanobacterium—that
cell that engulfed an alpha proteobacterium, thereby setting then evolved into plastids. According to the hypothesis illus-
the stage for the origin of eukaryotes. For example, research- trated in Figure 28.3, this plastid-bearing lineage gave rise to
ers recently reported the discovery of a new group of archaea, two lineages of photosynthetic protists, or algae: red algae
the lokiarchaeotes. In phylogenomic analyses, this group was and green algae.
identified as the sister group of the eukaryotes and its genome Let’s examine some of the steps in Figure 28.3 more
was found to encode many eukaryote-specific features. Was closely. First, recall that cyanobacteria are gram-negative
the host cell that engulfed an alpha proteobacterium a lokiar- and that gram-negative bacteria have two cell membranes,
chaeote? While this could be correct, it is also possible that the an inner plasma membrane and an outer membrane
host belonged to a different lineage of archaea or was closely that is part of the cell wall (see Figure 27.3). Plastids in
related to the archaea (but was not itself an archaean). In any red algae and green algae are also surrounded by two
case, current evidence indicates that the host was a relatively membranes. Transport proteins in these membranes are
complex cell in which certain features of eukaryotic cells had homologous to proteins in the inner and outer membranes
evolved, such as a cytoskeleton that enabled it to change of cyanobacteria, providing further support for the
shape (and thereby engulf the alpha proteobacterium). hypothesis that plastids originated from a cyanobacterial
endosymbiont.
Plastid Evolution: A Closer Look On several occasions during eukaryotic evolution, red algae
As you’ve seen, current evidence indicates that mitochon- and green algae underwent secondary endosymbiosis,
dria are descended from a bacterium that was engulfed by meaning they were ingested in the food vacuoles of hetero-
a host cell that was an archaean (or a close relative of the trophic eukaryotes and became endosymbionts themselves.
archaea). This event gave rise to the eukaryotes. There is For example, protists known as chlorarachniophytes likely
also much evidence that later in eukaryotic history, a lin- evolved when a heterotrophic eukaryote engulfed a green
eage of heterotrophic eukaryotes acquired an additional alga. Evidence for this process can be found within the

. Figure 28.3 Diversity of plastids produced by endosymbiosis. Studies of

plastid-bearing eukaryotes suggest that plastids evolved from a cyanobacterium that
was engulfed by an ancestral heterotrophic eukaryote (primary endosymbiosis). Plastid
That ancestor then diversified into red algae and green
algae, some of which were subsequently engulfed by
other eukaryotes (secondary endosymbiosis).

Stramenopiles
Secondary
Membranes endosymbiosis
are represented
as dark lines in
the cell.
Red alga
Cyanobacterium
1 2
3

Primary
endosymbiosis
Alveolates

Secondary Plastid
Nucleus
Heterotrophic endosymbiosis
eukaryote One of these
membranes was
lost in red and
green algal Euglenids
descendants.
Secondary
endosymbiosis
Green alga
VISUAL SKILLS Based on this diagram, which of
the following groups are likely to be more closely
related: stramenopiles and alveolates, or euglenids
Chlorarachniophytes
and chlorarachniophytes? Explain.

596 UNIT FIVE The Evolutionary History of Biological Diversity

c Figure 28.4 Inner plastid . Figure 28.6 A haptophyte and a cryptophyte. Although they
Nucleomorph membrane are major groups of protists, haptophytes and cryptophytes are not
within a plastid of a shown in Figure 28.5 because their phylogenetic relationships have
chlorarachniophyte. Outer plastid yet to be resolved. (a) The haptophyte Emiliania huxleyi is a key
membrane producer in the ocean. (b) The outer surface of this photosynthetic
cryptophyte is covered with protective plates. (SEM)

(a) (b)
Nuclear
pore-like gap

Nucleomorph

Thylakoid
membranes

2 om 8 om
engulfed cell, which contains a tiny vestigial nucleus, called
a nucleomorph (Figure 28.4). Genes from the nucleomorph are
still transcribed, and their DNA sequences indicate that the you read this chapter, it may be helpful to focus less on the
engulfed cell was a green alga. specific names of groups of organisms and more on why
the organisms are important and how ongoing research is
elucidating their evolutionary relationships.
Four Supergroups of Eukaryotes
Our understanding of the evolutionary history of eukary-
otic diversity has been in flux in recent years. Not only has CONCEPT CHECK 28.1

kingdom Protista been abandoned, but other hypotheses 1. Cite at least four examples of structural and functional
have been discarded as well. For example, many biologists diversity among protists.
once thought that the first lineage to have diverged from 2. Summarize the role of endosymbiosis in eukaryotic
evolution.
all other eukaryotes was the amitochondriate protists, organ-
3. MAKE CONNECTIONS After studying Figure 28.3, predict
isms without conventional mitochondria and with fewer how many distinct genomes are contained within the cell of
membrane-enclosed organelles than other protist groups. a chlorarachniophyte. Explain. (See Figures 6.17 and 6.18.)
But recent structural and DNA data have undermined For suggested answers, see Appendix A.
this hypothesis. Many of the so-called amitochondriate
protists have been shown to have mitochondria—though
reduced ones—and some of these organisms are now clas-
sified in distantly related groups. CONCEPT 28.2
The ongoing changes in our understanding of the phy-
logeny of protists pose challenges to students and instruc- Excavates include protists with
tors alike. Hypotheses about these relationships are a focus modified mitochondria and
of scientific activity, changing rapidly as new data cause
previous ideas to be modified or discarded. We’ll focus protists with unique flagella
here on one current hypothesis: the four supergroups of
Diplomonads
Excavata

eukaryotes shown in Figure 28.5. Because the root of the

Parabasalids
eukaryotic tree is not known, all four supergroups are
Euglenozoans
shown as diverging simultaneously from a common ances-
tor. We know that this is not correct, but we do not know SAR
Archaeplastida
which supergroup was the first to diverge from the other
Unikonta
three. In addition, major new groups of protists have been
discovered in recent years, but their relationship to the Now that we have examined some of the broad patterns in
supergroups shown in Figure 28.5 has yet to be resolved. eukaryotic evolution, we will look more closely at the four
Examples of such unresolved groups include haptophytes main groups of protists shown in Figure 28.5.
and cryptophytes (Figure 28.6), as well as the hemimas- We begin with Excavata (the excavates), a clade that was
tigophores, a novel group described in 2018. Overall, as originally proposed based on morphological studies of the

CHAPTER 28 Protists 597

. Figure 28.5 Exploring Protistan Diversity

The tree below represents a phylogenetic hypothesis for the relationships among eukaryotes on
Earth today. The eukaryotic groups at the branch tips are related in larger “supergroups,” labeled
vertically at the far right of the tree. Groups that were formerly classified in the kingdom Protista
are highlighted in yellow. Dotted lines indicate evolutionary relationships that are uncertain. For
clarity, this tree only includes representative clades from each supergroup. In addition, the recent
discoveries of many new groups of eukaryotes indicate that eukaryotic diversity is actually much
greater than shown here.

Diplomonads
Excavata

Excavata
Parabasalids Some members of this supergroup (whose
monophyly has been called into question
Euglenozoans by some recent genomic studies) have an
“excavated” groove on one side of
the cell body. Two major clades (the
parabasalids and diplomonads) have
Diatoms
Stramenopiles

highly reduced mitochondria; members

of a third clade (the euglenozoans) have
Brown algae flagella that differ in structure from
those of other organisms. Excavates
Oomycetes
include parasites such as Giardia, as well
as many predatory and photosynthetic
Dinoflagellates
species.
Alveolates

5 om

SAR
Apicomplexans

Ciliates

Radiolarians
Rhizarians

Forams

Cercozoans

Red algae
Archaeplastida

Chlorophytes
Green
algae

Charophytes

Plants
Giardia intestinalis, a diplomonad
parasite. This diplomonad (colorized
Amoebozoans

Tubulinids SEM), which lacks the characteristic surface

groove of the Excavata, inhabits the
Slime molds intestines of mammals. It can infect people
when they drink water contaminated with
Entamoebas feces containing Giardia cysts. Drinking
such water—even from a seemingly
Unikonta

Nucleariids pristine stream—can cause severe diarrhea.

Opisthokonts

Boiling the water kills the parasite.

Fungi

Choanoflagellates

Animals

DRAW IT Draw a simplified version of this phylogenetic tree that shows

only the four supergroups of eukaryotes. Now sketch how the tree would
look if the unikonts were the sister group to all other eukaryotes.

598 UNIT FIVE The Evolutionary History of Biological Diversity

 SAR Archaeplastida
This supergroup contains (and is named after) three large and This supergroup of eukaryotes includes red algae and green algae,
very diverse clades: Stramenopila, Alveolata, and Rhizaria. Stra- along with plants. Red algae and green algae include unicellular
menopiles include some of the most important photosynthetic or- species, colonial species, and multicellular species (including the
ganisms on Earth, such as the diatoms shown here. Alveolates also green alga Volvox). Many of the large algae known informally
include photosynthetic species, as well as important pathogens, as “seaweeds” are multicellular red or green algae. Protists in
such as Plasmodium, which causes malaria. According to one Archaeplastida include key photosynthetic species that form the
current hypothesis, stramenopiles and alveolates originated by base of the food web in many aquatic communities.
20 om
secondary endosymbiosis when a heterotrophic protist engulfed
25 om
a red alga.

50 om

Volvox, a multicellular freshwater green alga. This alga has two

types of differentiated cells, and so it is considered multicellular rather
than colonial. It resembles a hollow ball whose wall is composed of
Diatom diversity. These beautiful single-celled protists are hundreds of biflagellated cells (see inset LM) embedded in a gelatinous
important photosynthetic organisms in aquatic communities (LM). extracellular matrix; if isolated, these cells cannot reproduce. However,
the alga also contains cells that are specialized for either sexual or
The rhizarian subgroup of SAR includes many species of amoe- asexual reproduction. The large algae shown here will eventually
bas, most of which have pseudopodia that are threadlike in shape. release the small “daughter” algae that can be seen within them (LM).
Pseudopodia are extensions that can
Mastering Biology Video: Volvox
bulge from any portion of the cell;
they are used in movement and in Unikonta
the capture of prey.
This supergroup of eukaryotes, also called Amorphea, includes
amoebas that have lobe- or tube-shaped pseudopodia, as well as
animals, fungi, and non-amoeba protists that are closely related

100 om
to animals or fungi. According to one current hypothesis, the
unikonts were the first eukaryotic supergroup to diverge from
all other eukaryotes; however, this hypothesis has yet to be
widely accepted.

100 μm

Globigerina, a rhizarian in SAR. This species is a foram, a group

A unikont amoeba. This amoeba, the tubulinid Amoeba
whose members have threadlike pseudopodia that extend through
proteus, is using its pseudopodia to move.
pores in the shell, or test (LM). The inset shows a foram test, which is
hardened by calcium carbonate. Mastering Biology Video: Amoeba Pseudopodium

CHAPTER 28 Protists 599

cytoskeleton. Some members of this diverse group also have on the vaginal lining, promoting infection. Studies sug-
an “excavated” feeding groove on one side of the cell body. gest that T. vaginalis acquired this gene by horizontal gene
The excavates include the diplomonads, parabasalids, and transfer from bacterial parasites in the vagina. (Trichomonas
euglenozoans. Molecular data indicate that each of these infections also can occur in the urethra of males, though
three groups is monophyletic. In contrast, some recent often without symptoms.)
genomic studies indicate that the excavates are not mono-
phyletic, making Excavata the most controversial of our
four supergroups.
Euglenozoans
Protists called euglenozoans belong to a diverse clade that
includes predatory heterotrophs, photosynthetic autotrophs,
Diplomonads and Parabasalids mixotrophs, and parasites. The main morphological feature
The protists in these two groups lack plastids and have highly that distinguishes protists in this clade is the presence of a
reduced mitochondria (until recently, they were thought to rod with either a spiral or a crystalline structure inside each
lack mitochondria altogether). Most diplomonads and para- of their flagella (Figure 28.8). The two best-studied groups of
basalids are found in anaerobic environments. euglenozoans are the kinetoplastids and the euglenids.
Diplomonads have reduced mitochondria called
mitosomes. These organelles lack functional electron transport Kinetoplastids
chains and hence cannot use oxygen to help extract energy Protists called kinetoplastids have a single, large mito-
from carbohydrates and other organic molecules. Instead, chondrion that contains an organized mass of DNA called a
diplomonads get the energy they need from anaerobic kinetoplast. These protists include species that feed on prokary-
biochemical pathways. Many diplomonads are parasites, otes in freshwater, marine, and moist terrestrial ecosystems,
including the infamous Giardia intestinalis (see Figure 28.5), as well as species that parasitize animals, plants, and other
which inhabits the intestines of mammals. protists. For example, kinetoplastids in the genus Trypanosoma
Structurally, diplomonads have two equal-sized nuclei and infect humans and cause sleeping sickness (Figure 28.9), a
multiple flagella. Recall that eukaryotic flagella are exten- neurological disease that is invariably fatal if not treated. This
sions of the cytoplasm, consisting of bundles of microtubules disease currently afflicts about 10,000 people each year, mostly
covered by the cell’s plasma membrane (see Figure 6.24). in rural areas of Africa. The infection occurs via the bite of a
They are quite different from prokaryotic flagella, which are vector (carrier) organism, the African tsetse fly. Trypanosomes
filaments composed of globular proteins attached to the cell also cause Chagas’ disease, which is transmitted by bloodsuck-
surface (see Figure 27.7). ing insects and can lead to congestive heart failure.
Parabasalids also have reduced mitochondria; called Trypanosomes evade immune responses with an effec-
hydrogenosomes, these organelles generate some energy tive “bait-and-switch” defense. The surface of a trypano-
anaerobically, releasing hydrogen gas as a by-product. The some is coated with millions of copies of a single protein.
best-known parabasalid is Trichomonas vaginalis, a sexually However, before the host’s immune system can recognize
transmitted parasite that infects about 140 million people the protein and mount an attack, new generations of the
each year worldwide. T. vaginalis travels along the mucus-
coated lining of the human reproductive and urinary tracts . Figure 28.8 Euglenozoan flagellum. Most euglenozoans have a
by moving its flagella and by undulating part of its plasma crystalline rod inside one of their flagella (the TEM is a flagellum shown
membrane (Figure 28.7). In females, if the vagina’s normal in cross section). The rod lies alongside the 9 + 2 ring of microtubules
found in all eukaryotic flagella (compare with Figure 6.24).
acidity is disturbed, T. vaginalis can outcompete benefi-
cial microorganisms there and infect the vagina. One of

0.2 om
T. vaginalis’s genes encodes a product that allows it to feed Flagella

. Figure 28.7 The parabasalid parasite Trichomonas vaginalis.

(colorized SEM)

Flagella

8 om Crystalline rod
(cross section)

5 om
Undulating Ring of microtubules
membrane (cross section)

600 UNIT FIVE The Evolutionary History of Biological Diversity

. Figure 28.9 Trypanosoma, the kinetoplastid that causes CONCEPT CHECK 28.2
sleeping sickness. (colorized SEM)
1. Why do some biologists describe the mitochondria of
diplomonads and parabasalids as “highly reduced”?
2. WHAT IF? DNA sequence data for a diplomonad, a euglenid,
a plant, and an unidentified protist suggest that the unidenti-
fied species is most closely related to the diplomonad. Further
studies reveal that the unknown species has fully functional
mitochondria. Based on these data, at what point on the phy-
logenetic tree in Figure 28.5 did the mystery protist’s lineage
probably diverge from other eukaryote lineages? Explain.
Trypanosome For suggested answers, see Appendix A.

CONCEPT 28.3

Red blood
SAR is a highly diverse group
cell
of protists defined by DNA
9 om
similarities
Excavata
Diatoms
Brown algae Stramenopiles
parasite switch to another surface protein with a differ-
Oomycetes
ent molecular structure. Frequent changes in the surface Dinoflagellates
protein prevent the host from developing immunity. (The

SAR
Apicomplexans Alveolates
Scientific Skills Exercise in Chapter 43 explores this topic Ciliates

further.) About a third of Trypanosoma’s genome is dedi- Radiolarians

Forams Rhizarians
cated to producing these surface proteins. Cercozoans
Archaeplastida
Unikonta
Euglenids
A euglenid has a pocket at one end of the cell from which Our second supergroup, referred to as SAR, was proposed
one or two flagella emerge (Figure 28.10). Some euglenids are recently based on whole-genome DNA sequence analyses.
mixotrophs: They perform photosynthesis when sunlight is These studies found that three major clades of protists—the
available, but when it is not, they can become heterotrophic, stramenopiles, alveolates, and rhizarians—form a monophy-
absorbing organic nutrients from their environment. Many letic supergroup. This supergroup contains a large, extremely
other euglenids engulf prey by phagocytosis. diverse collection of protists. To date, this supergroup has not

▶ Figure 28.10 Euglena, a euglenid Long flagellum

commonly found in pond water.
Mastering Biology Video: Euglena
Eyespot: pigmented
organelle that
functions as a light
shield, allowing light
from only a certain Light detector:
direction to strike swelling near the base
the light detector of the long flagellum;
detects light that is
not blocked by the
Short flagellum eyespot. As a result,
Euglena moves
Contractile vacuole toward light of
appropriate intensity,
Nucleus an important
adaptation that
Chloroplast enhances
photosynthesis.
Plasma membrane
5 om Pellicle: protein bands beneath
Euglena (LM) the plasma membrane that
provide strength and flexibility

CHAPTER 28 Protists 601

received a formal name but is instead known by the first let- c Figure 28.12 The
ters of its major clades: SAR. diatom Triceratium
morlandii. (colorized SEM)
Some morphological and DNA sequence data suggest that
two of these groups, the stramenopiles and alveolates, origi-
nated more than a billion years ago, when a common ancestor the surface of the sea may
of these two clades engulfed a single-celled, photosynthetic contain millions of these
red alga. Because red algae are thought to have originated by microscopic algae. The
primary endosymbiosis (see Figure 28.3), such an origin for abundance of diatoms
the stramenopiles and alveolates is referred to as secondary in the past is also evident
endosymbiosis. Others question this idea, noting that some
40 om
in the fossil record, where
species in these groups lack plastids or their remnants (includ- massive accumulations of
ing any trace of plastid genes in their nuclear DNA). fossilized diatom walls are
Despite its lack of a formal name, the SAR supergroup rep- major constituents of sediments known as diatomaceous earth.
resents the best current hypothesis for the phylogeny of the These sediments are mined for their quality as a filtering
three large protist clades to which we now turn. medium and for many other uses.
Diatoms are so widespread and abundant that their pho-
Stramenopiles tosynthetic activity affects global carbon dioxide (CO2) lev-
els. Diatoms have this effect in part because of events that
One major subgroup of SAR, the stramenopiles, includes
occur during episodes of rapid population growth, or blooms,
some of the most important photosynthetic organisms on the
when ample nutrients are available. Typically, diatoms are
planet. Their name (from the Latin stramen, straw, and pilos,
eaten by a variety of protists and invertebrates, but during a
hair) refers to their characteristic flagellum, which has numer-
bloom, many escape this fate. When these uneaten diatoms
ous fine, hairlike projections. In most stramenopiles, this
die, their bodies sink to the ocean floor. It takes decades, or
“hairy” flagellum is paired with a shorter “smooth” (nonhairy)
even centuries, for dead diatoms that sink to the ocean floor
flagellum (Figure 28.11). Here we’ll focus on three groups of
to be broken down by bacteria and other decomposers. As a
stramenopiles: diatoms, oomycetes, and brown algae.
result, the carbon in their bodies remains there for some time,
rather than being released immediately as CO2 as the decom-
. Figure 28.11 Stramenopile flagella. Most stramenopiles, such posers respire. The overall effect of these events is that CO2
as Synura petersenii, have two flagella: one covered with fine, stiff absorbed by diatoms during photosynthesis is transported, or
hairs and a shorter one that is smooth.
“pumped,” to the ocean floor.
With an eye toward reducing global warming by lowering
atmospheric CO2 levels, some scientists advocate promoting
diatom blooms by fertilizing the ocean with essential
nutrients such as iron. In one recent study, researchers
Hairy found that CO2 was indeed pumped to the ocean floor
flagellum
after iron was added to a small region of the ocean. Further
Smooth

2.5 om
flagellum tests are planned to examine whether iron fertilization has
undesirable side effects (such as oxygen depletion or the
production of nitrous oxide, a more potent greenhouse
gas than CO2).

Diatoms Brown Algae

A key group of photosynthetic protists, diatoms are unicel- The largest and most complex algae are brown algae. All are
lular algae that have a unique glass-like wall made of silicon multicellular, and most are marine. Brown algae are especially
dioxide embedded in an organic matrix (Figure 28.12). The common along temperate coasts that have cold-water cur-
wall consists of two parts that overlap like a shoe box and its rents. They owe their characteristic brown or olive color to
lid. These walls provide effective protection from the crush- the carotenoids in their plastids.
ing jaws of predators: Live diatoms can withstand pressures as Many of the species commonly called “seaweeds” are brown
great as 1.4 million kg/m2, equal to the pressure under each algae. Some brown algal seaweeds have specialized structures
leg of a table supporting an elephant! that resemble organs in plants, such as a rootlike holdfast,
With an estimated 100,000 living species, diatoms are a which anchors the alga, and a stemlike stipe, which supports
highly diverse group of protists (see Figure 28.5). They are the leaflike blades (Figure 28.13). Unlike plants, however,
among the most abundant photosynthetic organisms both brown algae lack true tissues and organs. Moreover, morpho-
in the ocean and in lakes: One bucket of water scooped from logical and DNA data show that these similarities evolved

602 UNIT FIVE The Evolutionary History of Biological Diversity

. Figure 28.13 Seaweeds: adapted to life at the ocean’s independently in the algal and plant lineages and are thus
margins. The sea palm (Postelsia) lives on rocks along the coast analogous, not homologous. In addition, while plants have
of the northwestern United States and western Canada. The body
adaptations (such as rigid stems) that provide support against
of this brown alga is well adapted to maintaining a firm foothold
despite the crashing surf. gravity, brown algae have adaptations that enable their main
photosynthetic surfaces (the leaflike blades) to be near the
water surface. Some brown algae accomplish this task with gas-
filled, bubble-shaped floats. Giant brown algae known as kelps
that live in deep waters have such floats in their blades, which
are attached to stipes that can rise as much as 60 m from the
Blade
seafloor—more than half the length of a football field.
Brown algae are important commodities for humans. Some
species are eaten, such as Laminaria (Japanese “kombu”),
which is used in soups. In addition, the cell walls of brown
Stipe
algae contain a gel-forming substance, called algin, which is
used to thicken many processed foods, including pudding
and salad dressing.

Alternation of Generations
Holdfast 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, as shown in Figure 28.14 for the brown alga

. Figure 28.14 The life cycle of the brown alga Laminaria:

an example of alternation of generations. 1 The sporophytes are usually Key
found in water just below the
line of the lowest tides, attached Haploid (n)
to rocks by branching holdfasts.
Diploid (2n)

2 Cells on the surface

Sporangia of the blade develop
into sporangia.

3 Sporangia produce
zoospores by meiosis.
MEIOSIS
Sporophyte
(2n)
Zoospore
4 The zoospores are all
structurally alike, but
about half of them divide
asexually (by mitosis)
7 The zygotes
and develop into male
grow into new Female gametophytes; the other
sporophytes
Developing half divide asexually
while attached
sporophyte and develop into
to the remains
Gametophytes female gametophytes.
of the female
gametophyte. (n)
Zygote
(2n)
Male

Egg
Mature female FERTILIZATION
gametophyte
5 Male gametophytes release sperm,
(n) and female gametophytes produce eggs,
Sperm which remain attached to the female
gametophyte. Eggs secrete a chemical
VISUAL SKILLS Based on this diagram, are the signal that attracts sperm of the same
sperm shown in 5 genetically identical to one 6 Sperm fertilize species, thereby increasing the
another? Explain. the eggs. probability of fertilization in the ocean.

CHAPTER 28 Protists 603

Laminaria. Although haploid and diploid conditions alternate are key differences between oomycetes and fungi. Among
in all sexual life cycles—human gametes, for example, are them, oomycetes typically have cell walls made of cellulose,
haploid—the term alternation of generations applies only to life whereas the walls of fungi consist mainly of another poly-
cycles in which both haploid and diploid stages are multicel- saccharide, chitin. Data from molecular systematics have
lular. As you will read in Concept 29.1, alternation of genera- confirmed that oomycetes are not closely related to fungi.
tions also evolved in plants. In both oomycetes and fungi, the high surface-to-volume
As detailed in Figure 28.14, the diploid individual in ratio of filamentous structures enhances the uptake of
this complex algal life cycle is called the sporophyte because nutrients from the environment.
it produces spores. The spores are haploid and move by Although oomycetes descended from plastid-bearing
means of flagella; they are called zoospores. The zoospores ancestors, they no longer have plastids and do not perform
divide by mitosis and develop into haploid, multicellular photosynthesis. Instead, they typically acquire nutrients as
male and female gametophytes, which produce gametes. decomposers or parasites. Most water molds are decompos-
The union of two gametes (fertilization) results in a diploid ers that grow as cottony masses on dead algae and animals,
zygote, which matures and gives rise to a new multicellular mainly in freshwater habitats (Figure 28.15b). White rusts
sporophyte. and downy mildews generally live on land as plant parasites.
In Laminaria, the two generations are heteromorphic, The ecological impact of oomycetes can be significant.
meaning that the sporophytes and gametophytes are struc- For example, the oomycete Phytophthora infestans causes
turally different. Other algal life cycles have an alternation potato late blight, which turns the stalks and stems of
of isomorphic generations, in which the sporophytes and potato (and tomato) plants into black slime. Late blight
gametophytes look similar to each other, although they differ contributed to the devastating Irish famine of the 19th cen-
in chromosome number. tury, in which a million people died and at least that many
were forced to leave Ireland. The disease remains a major
Oomycetes (Water Molds and Their Relatives) problem today, causing crop losses as high as 70% in some
Oomycetes include the water molds, the white rusts, regions. Globally, about $1 billion is spent each year on pes-
and the downy mildews. Based on their morphology, ticides to control the parasite.
these organisms were previously classified as fungi (in
fact, oomycete means “egg fungus”). For example, many
oomycetes have multinucleate filaments (hyphae) that Alveolates
resemble fungal hyphae (Figure 28.15a). However, there Members of the next subgroup of SAR, the alveolates, have
membrane-enclosed sacs (alveoli) just under the plasma
. Figure 28.15 Oomycetes. membrane (Figure 28.16). Alveolates are abundant in many
habitats and include a wide range of photosynthetic and het-
100 om
(a) Closeup of oomycete
hyphae (LM)
erotrophic protists. We’ll discuss three alveolate clades here:
a group of flagellates (the dinoflagellates), a group of parasites
(the apicomplexans), and a group of protists that move using
cilia (the ciliates).

. Figure 28.16 Alveoli. These sacs under the plasma membrane

are a characteristic that distinguishes alveolates from other
eukaryotes (TEM).

Flagellum Alveoli

(b) Water mold hyphae

growing on a goldfish

Alveolate
0.2 om

604 UNIT FIVE The Evolutionary History of Biological Diversity

. Figure 28.17 Dinoflagellates. blooms occur, toxins produced by certain dinoflagellates
(a) Dinoflagellate flagella. have caused massive kills of invertebrates and fishes. Humans
Beating of the spiral who eat molluscs that have accumulated the toxins are
flagellum, which lies in a
affected as well, sometimes fatally. A 2017 study found that
groove that encircles the
cell, makes this specimen ocean warming (caused by ongoing climate change) has facil-
of Pfiesteria shumwayae itated more frequent blooms of toxic dinoflagellates, making
spin (colorized SEM).
them a growing threat to human health.

Apicomplexans
Flagella
Nearly all apicomplexans are parasites of animals—and
virtually all animal species examined so far are attacked
by these parasites. The parasites spread through their host
as tiny infectious cells called sporozoites. Apicomplexans
are so named because one end (the apex) of the sporozoite

4 om
cell contains a complex of organelles specialized for
penetrating host cells and tissues. Although apicomplexans
are not photosynthetic, recent data show that they
retain a modified plastid (apicoplast), most likely of red
algal origin.
Most apicomplexans have intricate life cycles with both
sexual and asexual stages. Those life cycles often require
two or more host species for completion. For example,
Plasmodium, the parasite that causes malaria, lives in both
mosquitoes and humans (Figure 28.18).
(b) Red tide in the Gulf Historically, malaria has rivaled tuberculosis as the
of Carpentaria in leading cause of human death by infectious disease. The
northern Australia. incidence of malaria was diminished in the 1960s by
The red color is due to
high concentrations of insecticides that reduced carrier populations of Anopheles
a carotenoid-containing mosquitoes and by drugs that killed Plasmodium in
dinoflagellate.
humans. But the emergence of resistant varieties of both
Anopheles and Plasmodium has led to a resurgence of
Mastering Biology Video: Dinoflagellate malaria. About 220 million people in the tropics are cur-
rently infected, and 450,000 die each year. In regions
where malaria is common, the lethal effects of this disease
Dinoflagellates have resulted in the evolution of high frequencies of the
The cells of many dinoflagellates are reinforced by cellu- sickle-cell allele; for an explanation of this connection,
lose plates. Grooves in this “armor” house two flagella, one of see Figure 23.18.
which causes dinoflagellates (from the Greek dinos, whirling) The search for malarial vaccines has been hampered by
to spin as they move through the waters of their marine and the fact that Plasmodium lives mainly inside cells, hidden
freshwater communities (Figure 28.17a). from the host’s immune system. And, like trypanosomes,
Although their ancestors may have originated by second- Plasmodium continually changes its surface proteins. Even
ary endosymbiosis (see Figure 28.3), roughly half of all so, significant progress was made when European regulators
dinoflagellate species are now purely heterotrophic. Others recently approved the world’s first licensed malarial vaccine.
are important species of phytoplankton (photosynthetic In 2019, routine use of this vaccine began in selected regions
plankton, which include photosynthetic bacteria as well of Africa. However, this vaccine, which targets a protein on
as algae); many photosynthetic dinoflagellates are the surface of sporozoites, provides only partial protection
mixotrophic. against malaria. As a result, researchers continue to study
Periods of explosive population growth (blooms) in dino- other potential vaccine targets, including the apicoplast.
flagellates sometimes cause a phenomenon called “red tide” This approach may be effective because the apicoplast is a
(Figure 28.17b). The blooms make coastal waters appear modified plastid; as such, it descended from a cyanobacte-
brownish red or pink because of the presence of carotenoids, rium and hence has different metabolic pathways from those
the most common pigments in dinoflagellate plastids. When in human patients.

CHAPTER 28 Protists 605

. Figure 28.18 The two-host life cycle of 2 The sporozoites enter the person’s
Plasmodium, the apicomplexan that causes 1 An infected Anopheles liver cells. After several days, the sporozoites
malaria. mosquito bites a person, undergo multiple mitotic divisions and become
injecting Plasmodium merozoites, which use their apical complex
? Are morphological differences between sporozoites in its saliva. to penetrate red blood cells (see TEM below).
sporozoites, merozoites, and gametocytes
caused by different genomes
0.5 om
or by differences in gene
expression? Explain. Inside mosquito Inside human Merozoite

Mastering Biology
Animation: Life Cycle Sporozoites
of a Malaria Parasite (n)
Liver

8 An oocyst develops
from the zygote in the wall
of the mosquito’s gut. The Liver cell
oocyst releases thousands
of sporozoites, which
migrate to the mosquito’s Oocyst
salivary gland.
Red blood Apex
MEIOSIS
Merozoite cell
(n)
Zygote
(2n) Red blood 3 The merozoites divide
cells asexually inside the red
blood cells. At intervals of
48 or 72 hours (depending
on the species), large
numbers of merozoites
7 Fertilization occurs break out of the blood
in the mosquito’s FERTILIZATION cells, causing periodic chills
digestive tract, and a and fever. Some of the
zygote forms. merozoites infect other
red blood cells.
Gametes
Gametocytes
Key (n)

Haploid (n) 4 Some merozoites

form gametocytes.
Diploid (2n)

6 Gametes form from gametocytes; each male 5 Another Anopheles mosquito bites the infected person
gametocyte produces several slender male gametes. and picks up Plasmodium gametocytes along with blood.

Ciliates (Figure 28.19b). Ciliates generally reproduce asexually by

The ciliates are a large and varied group of protists named for binary fission, during which the existing macronucleus dis-
their use of cilia to move and feed (Figure 28.19a). Most ciliates integrates and a new one is formed from the cell’s micronu-
are predators, typically of bacteria or of other protists. Their cilia clei. Each macronucleus typically contains multiple copies
may completely cover the cell surface or may be clustered in a few of the ciliate’s genome. Genes in the macronucleus control
rows or tufts. In certain species, rows of tightly packed cilia func- the everyday functions of the cell, such as feeding, waste
tion collectively in locomotion. Other ciliates scurry about on removal, and maintaining water balance.
leg-like structures constructed from many cilia bonded together. Mastering Biology Video: Ciliate Movement in Stentor
A distinctive feature of ciliates is the presence of two types
of nuclei: tiny micronuclei and large macronuclei. A cell has
one or more nuclei of each type. Genetic variation results Rhizarians
from conjugation, a sexual process in which two individu- Our next subgroup of SAR is the rhizarians. Many species
als exchange haploid micronuclei but do not reproduce in this group are amoebas, protists that move and feed by

606 UNIT FIVE The Evolutionary History of Biological Diversity

. Figure 28.19 Structure and function in the ciliate Paramecium caudatum.

Paramecium constantly takes in water Contractile Cilia along a funnel-shaped oral groove
by osmosis from its hypotonic vacuole move food (mainly bacteria) into the
environment. Bladderlike contractile cell mouth, where the food is engulfed
vacuoles accumulate excess water from into food vacuoles by phagocytosis.
radial canals and periodically expel it
through the plasma membrane.
Oral groove

50 om
Thousands of cilia cover the Cell mouth
surface of Paramecium.

Food vacuoles fuse with lysosomes (not

shown). As the food is digested, the
Micronucleus vacuoles follow a looping path through
the cell. Wastes are released when the
Macronucleus vacuoles fuse with a specialized region
of the plasma membrane that functions
as an anal pore.

(a) Feeding, waste removal, and water balance

1 Two cells of compatible 2 Meiosis of micronuclei 3 Three micronuclei in each cell

mating strains align side produces four haploid disintegrate. The remaining micro-
by side and partially fuse. micronuclei in each cell. nucleus in each cell divides by mitosis.

MEIOSIS 4 The cells swap

one micronucleus.

Haploid
Diploid micronucleus
Compatible
micronucleus
mates
The original Diploid
macronucleus micronucleus
disintegrates.

MICRONUCLEAR
FUSION
5 The cells
separate.

Key
9 Two rounds of
binary fission yield 8 Four micro- 7 Three rounds 6 The two Conjugation
four daughter cells. nuclei become of mitosis produce micronuclei fuse.
macronuclei. eight micronuclei. Asexual
reproduction
(b) Conjugation and reproduction

MAKE CONNECTIONS The events shown in steps 4 through 6 of this diagram have Mastering Biology Video: Paramecium
a similar overall effect to what event in the human life cycle (see Figure 13.5)? Explain.

means of pseudopodia, extensions that may bulge from across many distantly related eukaryotic taxa. Most amoe-
almost anywhere on the cell surface. An amoeba moves by bas that are rhizarians differ morphologically from other
extending a pseudopodium and anchoring its tip; more cyto- amoebas by having threadlike pseudopodia. Rhizarians also
plasm then streams into the pseudopodium. Amoebas do not include flagellated (non-amoeboid) protists that feed using
constitute a monophyletic group; instead, they are dispersed threadlike pseudopodia.

CHAPTER 28 Protists 607

 We’ll examine three groups of rhizarians here: radiolar- . Figure 28.21 Fossil forams. By measuring the magnesium
ians, forams, and cercozoans. content in fossilized forams like these, researchers seek to learn how
ocean temperatures have changed over time. Forams take up more
magnesium in warmer water than in colder water.
Radiolarians
The protists called radiolarians have delicate, intricately
symmetrical internal skeletons that are generally made
of silica. The pseudopodia of these mostly marine pro-
tists radiate from the central body (Figure 28.20) and are
reinforced by bundles of microtubules. The microtubules
are covered by a thin layer of cytoplasm, which engulfs
smaller microorganisms that become attached to the pseu-
dopodia. Cytoplasmic streaming then carries the captured
prey into the main part of the cell. After radiolarians die,
their skeletons settle to the seafloor, where they have
accumulated as an ooze that is hundreds of meters thick in
some locations.

. Figure 28.20 A radiolarian. Numerous threadlike pseudopodia

radiate from the central body of this radiolarian (LM).

remains of other protists, the fossilized tests of forams are part

of marine sediments, including sedimentary rocks that are
now land formations. Foram fossils are excellent markers for
correlating the ages of sedimentary rocks in different parts of
the world. Researchers are also studying these fossils to obtain
information about climate change and its effects on the
oceans and their life (Figure 28.21).

Cercozoans
First identified in molecular phylogenies, the cercozoans are
a large group of amoeboid and flagellated protists that feed
Pseudopodia using threadlike pseudopodia. Cercozoan protists are com-

200 om
mon inhabitants of marine, freshwater, and soil ecosystems.

. Figure 28.22 A second case of primary endosymbiosis?

The cercozoan Paulinella conducts photosynthesis in a unique
Forams sausage-shaped structure called a chromatophore (LM).
Chromatophore membranes include a peptidoglycan layer,
The protists called foraminiferans (from the Latin
indicating that they are derived from a bacterium. DNA evidence
foramen, little hole, and ferre, to bear), or forams, shows that chromatophores are derived from a different
are named for their porous shells, called tests (see cyanobacterium than that from which plastids are derived.
Figure 28.5). Foram tests consist of a single piece of organic
material that typically is hardened with calcium carbonate.
The pseudopodia that extend through the pores function Chromatophore
in swimming, test formation, and feeding. Many forams
also derive nourishment from the photosynthesis of sym-
biotic algae that live within the tests.
Forams are found in both the ocean and fresh water. Most
species live in sand or attach themselves to rocks or algae,
but some live as plankton. The largest forams, though single-
celled, have tests several centimeters in diameter.

5 om
Ninety percent of all identified species of forams are
known from fossils. Along with the calcium-containing

608 UNIT FIVE The Evolutionary History of Biological Diversity

 Most cercozoans are heterotrophs. Many are parasites of plants in Chapters 29 and 30; here we will look at the diver-
plants, animals, or other protists; many others are preda- sity of their closest algal relatives, red algae and green algae.
tors. The predators include the most important consumers
of bacteria in aquatic and soil ecosystems, along with spe- Red Algae
cies that eat other protists, fungi, and even small animals.
Many of the 6,000 known species of red algae (rhodo-
One small group of cercozoans, the chlorarachniophytes
phytes, from the Greek rhodos, red) are reddish, owing to the
(mentioned earlier in the discussion of secondary endosym-
photosynthetic pigment phycoerythrin, which masks the
biosis), are mixotrophic: These organisms ingest smaller
green of chlorophyll (Figure 28.23). However, other species
protists and bacteria as well as perform photosynthesis. At
(those adapted to shallow water) have less phycoerythrin. As
least one other cercozoan, Paulinella chromatophora, is an
a result, red algal species may be greenish red in very shallow
autotroph, deriving its energy from light and its carbon
water, bright red at moderate depths, and almost black in
from CO2. Paulinella (Figure 28.22) appears to represent an
deep water. Some species lack pigmentation altogether and
intriguing additional evolutionary example of a eukaryotic
live as heteroptrophic parasites on other red algae.
lineage that obtained its photosynthetic apparatus directly
from a cyanobacterium.
. Figure 28.23 Red algae.
CONCEPT CHECK 28.3
c Bonnemaisonia
1. Explain why forams have such a well-preserved fossil record. hamifera. This red alga
2. WHAT IF? Would you expect the plastid DNA of photo- has a filamentous form.
synthetic dinoflagellates and diatoms to be more similar
to the nuclear DNA of plants (domain Eukarya) or to the
chromosomal DNA of cyanobacteria (domain Bacteria)?
Explain.
3. MAKE CONNECTIONS Which of the three life cycles in
Figure 13.6 exhibits alternation of generations? How does it
differ from the other two?
4. MAKE CONNECTIONS Review Figures 9.2 and 10.5, and
then summarize how CO2 and O2 are both used and pro-
duced by chlorarachniophytes and other aerobic algae.
For suggested answers, see Appendix A.

CONCEPT 28.4 8 mm

Red algae and green algae are b Dulse (Palmaria palmata).

This edible species has a
the closest relatives of plants ”leafy” form.

Excavata . Nori. The red alga Porphyra is the source of

SAR a traditional Japanese food.
The seaweed is
Archaeplastida

Red algae
grown on nets in
Chlorophytes shallow coastal
Green algae
Charophytes waters.
Plants
Unikonta

As described earlier, morphological and molecular evidence

indicates that plastids arose when a heterotrophic protist
acquired a cyanobacterial endosymbiont. Later, photosyn-
thetic descendants of this ancient protist evolved into red
algae and green algae (see Figure 28.3), and the lineage that
produced green algae then gave rise to plants. Together, red
algae, green algae, and plants make up our third eukaryotic
supergroup, which is called Archaeplastida. Archaeplastida
Paper-thin, glossy sheets of dried nori
is a monophyletic group that descended from the ancient make a mineral-rich wrap for rice,
protist that engulfed a cyanobacterium. We will examine seafood, and vegetables in sushi.

CHAPTER 28 Protists 609

 Red algae are abundant in the warm coastal waters of tropi- . Figure 28.24 Examples of large chlorophytes.

10 om
cal oceans. Some of their photosynthetic pigments, including
(a) Pediastrum, a pond alga.
phycoerythrin, allow them to absorb blue and green light, Chlorophytes in this genus
which penetrate relatively far into the water. A species of red form daughter colonies with
alga has been discovered near the Bahamas at a depth of more the same number and
arrangement of cells as the
than 260 m. There are also a small number of freshwater and parent colony.
terrestrial species.
Most red algae are multicellular. Although none are as
big as the giant brown kelps, the largest multicellular red
algae are included in the informal designation “seaweeds.”
You may have eaten one of these multicellular red algae,
(b) Ulva, or sea
Porphyra (Japanese “nori”), as crispy sheets or as a wrap for 2 cm
lettuce. This
sushi (see Figure 28.23). Red algae reproduce sexually and multicellular, edible
have diverse life cycles in which alternation of generations chlorophyte has
differentiated
is common. However, unlike other algae, red algae do not structures, such as
have flagellated gametes, so they depend on water currents its leaflike blades
to bring gametes together for fertilization. and a rootlike
holdfast that
anchors the alga.

Green Algae
The grass-green chloroplasts of green algae have a struc-
ture and pigment composition much like the chloroplasts
of plants. Molecular systematics and cellular morphology
leave little doubt that green algae and plants are closely (c) Caulerpa, an
intertidal chloro-
related. In fact, some systematists now advocate including
phyte. The branched
green algae in an expanded “plant” kingdom, Viridiplantae filaments lack cross-
(from the Latin viridis, green). Phylogenetically, this walls and thus are
multinucleate. In
change makes sense, since otherwise the green algae are effect, the body of
a paraphyletic group. this alga is one huge
Green algae can be divided into two main groups, the cha- ”supercell.”
rophytes and the chlorophytes. The charophytes include the 2. The formation of true multicellular bodies by cell divi-
algae most closely related to plants, and we will discuss them sion and differentiation, as in Volvox (see Figure 28.5)
along with plants in Chapter 29. and Ulva (Figure 28.24b).
The second group, the chlorophytes (from the Greek
3. The repeated division of nuclei with no cytoplasmic
chloros, green), includes more than 7,000 species. Most live
division, as in Caulerpa (Figure 28.24c).
in fresh water, but there are also many marine and some ter-
restrial species. The simplest chlorophytes are unicellular Most chlorophytes have complex life cycles, with both
organisms such as Chlamydomonas, which resemble gametes sexual and asexual reproductive stages. Nearly all species of
of more complex chlorophytes. Various species of unicel- chlorophytes reproduce sexually by means of biflagellated
lular chlorophytes live independently in aquatic habitats as gametes that have cup-shaped chloroplasts (Figure 28.25).
phytoplankton or inhabit damp soil. Some live symbiotically Alternation of generations has evolved in some chlorophytes,
within other eukaryotes, contributing part of their photo- including Ulva.
synthetic output to the food supply of their hosts. Still other
chlorophytes live in environments exposed to intense vis- Mastering Biology Animation: Alternation of Generations
in a Protist
ible and ultraviolet radiation; these species are protected by
radiation-blocking compounds in their cytoplasm, cell wall,
or zygote coat. CONCEPT CHECK 28.4

Larger size and greater complexity evolved in green algae 1. Contrast red algae and brown algae.
by three different mechanisms: 2. Why is it accurate to say that Ulva is truly multicellular but
Caulerpa is not?
1. The formation of colonies of individual cells, as
3. WHAT IF? Suggest a possible reason why species in the
seen in Pediastrum (Figure 28.24a) and other species green algal lineage may have been more likely to colonize
that contribute to the stringy masses known as pond land than species in the red algal lineage.
scum. For suggested answers, see Appendix A.

610 UNIT FIVE The Evolutionary History of Biological Diversity

. Figure 28.25 The life cycle of Chlamydomonas, a unicellular chlorophyte.

Flagella 1 om 2 In response to a nutrient shortage,

drying of the enviroment, or other
stress, cells develop into gametes.
Cell wall
1 In Chlamydomonas,
mature cells are haploid
and contain a single 3 Gametes of different
cup-shaped chloroplast. mating types (designated
Nucleus
– + and –) fuse (fertilization),
forming a diploid zygote.

Cross +
section of Gamete
cup-shaped (n) –
+
chloroplast
(TEM)

7 These daughter cells develop Zoospore

Mature cell FERTILIZATION
flagella and cell walls and then emerge as
swimming zoospores from the parent cell. (n)
The zoospores develop into mature ASEXUAL
REPRODUCTION SEXUAL
haploid cells. REPRODUCTION Zygote
(2n)

–
+

Key 6 When a mature cell reproduces + – MEIOSIS

asexually, it resorbs its flagella and then
Haploid (n) undergoes two rounds of mitosis, forming
Diploid (2n) four cells (more in some species). 4 The zygote secretes
a durable coat that
protects the cell from
harsh conditions.
5 After a dormant period,
VISUAL SKILLS Circle the stage(s) in the diagram in which clones are meiosis produces four haploid
formed, producing additional new daughter cells that are genetically individuals (two of each mating
identical to the parent cell(s). type) that emerge and mature.

CONCEPT 28.5 groups). Each of these two major clades is strongly supported
by molecular systematics. Support for the close relationship
Unikonts include protists that between amoebozoans and opisthokonts is provided by com-
parisons of myosin proteins and by some (but not all) studies
are closely related to fungi and based on multiple genes or whole genomes.
animals An ongoing debate involving the unikonts concerns the
root of the eukaryotic tree. Recall that the root of a phylo-
Excavata
genetic tree anchors the tree in time: Branch points close to
SAR
Archaeplastida the root are the oldest. At present, the root of the eukary-
Tubulinids
Slime molds otic tree is uncertain; hence, we do not know which super-
Entamoebas group of eukaryotes was the first to diverge from all other
Unikonta

Nucleariids
Fungi eukaryotes. If the root of the eukaryotic tree were known,
Choanoflagellates it would help scientists infer characteristics of the common
Animals
ancestor of all eukaryotes.
Unikonta (also called Amorphea) is an extremely diverse In trying to determine the root of the eukaryotic tree,
supergroup of eukaryotes that includes animals, fungi, and researchers have based their phylogenies on different sets of
some protists. There are two major clades of unikonts, the genes, with conflicting results. Researchers have also tried a
amoebozoans (tubulinids and close protist relatives) and different approach, based on tracing the evolutionary history
the opisthokonts (animals, fungi, and closely related protist of genes that were transferred to eukaryotes from bacterial

CHAPTER 28 Protists 611

 . Figure 28.26 Inquiry Tubulinids
Tubulinids constitute a large and varied group of amoebozo-
What is the root of the eukaryotic tree?
ans that have lobe- or tube-shaped pseudopodia. These uni-
Experiment Responding to the difficulty in determining cellular protists are ubiquitous in soil as well as freshwater and
the root of the eukaryotic phylogenetic tree, Romain Derelle marine environments. Most are heterotrophs that actively
and colleagues used a new approach. They studied 39 nuclear seek and consume bacteria and other protists; one such tubu-
genes that eukaryotes acquired by horizontal gene transfer linid species, Amoeba proteus, is shown in Figure 28.5. Some
from a single bacterial source: the alpha proteobacterium
endosymbiont that gave rise to the mitochondrion. These 39 tubulinids also feed on detritus (nonliving organic matter).
genes were known to be orthologous (see Figure 26.18). As
a result, differences among these genes reflect the history Slime Molds
of speciation events that have occurred since the eukaryotes
began to diverge from their last common ancestor—making
Slime molds, or mycetozoans (from the Latin, meaning “fun-
these genes well suited for inferring the root of the eukary- gus animals”), once were thought to be fungi because, like
otic tree. Derelle and colleagues obtained the sequences of fungi, they produce fruiting bodies that aid in spore dispersal.
these genes for 67 species of eukaryotes representing a wide However, DNA sequence analyses indicate that the resem-
range of eukaryotic taxa. Those sequences were then used to
infer evolutionary relationships among the four eukaryotic blance between slime molds and fungi is a case of evolution-
supergroups shown in Figure 28.5. ary convergence. Slime molds have diverged into two main
Results branches, plasmodial slime molds and cellular slime molds.
Unikonta We’ll compare their characteristics and life cycles.

Plasmodial Slime Molds Many plasmodial slime molds

are brightly colored, often yellow or orange (Figure 28.27).
Excavata
As they grow, they form a mass called a plasmodium, which
can be many centimeters in diameter. (Don’t confuse a slime
Common ancestor
of all eukaryotes mold’s plasmodium with the genus Plasmodium, which
SAR
includes the parasitic apicomplexan that causes malaria.)
Despite its size, the plasmodium is not multicellular; it is a sin-
Archaeplastida
gle mass of cytoplasm that is undivided by plasma membranes
and that contains many nuclei. This “supercell” is the product
Conclusion The results show that Excavata, SAR, and of mitotic nuclear divisions that are not followed by cytokine-
Archaeplastida form a clade, which supports the hypothesis sis. The plasmodium extends pseudopodia through moist soil,
that the root of the tree is located between the unikonts and
leaf mulch, or rotting logs, engulfing food particles by phago-
all other eukaryotes.
cytosis as it grows. If the habitat begins to dry up or there is no
Data from R. Derelle et al., Bacterial proteins pinpoint a single eukaryotic root,
Proceedings of the National Academy of Sciences USA 112:E693–699 (2015). food left, the plasmodium stops growing and differentiates
WHAT IF? Other recent evidence indicates that the root of the into fruiting bodies that function in sexual reproduction.
eukaryotic tree may lie between a clade that includes unikonts and
excavates and all other eukaryotes. Draw the tree suggested by this c Figure 28.27
result. A mature
plasmodium, the
feeding stage in
the life cycle of a
plasmodial slime
donors (Figure 28.26). Results from this approach indicate mold.

that Excavata, SAR, and Archaeplastida share a more recent

common ancestor than any of them does with Unikonta.
This suggests that the root of the tree is located between the
4 cm
unikonts and all other eukaryotes, which implies that the
unikonts were the first eukaryotic supergroup to diverge from
all other eukaryotes. This idea remains controversial and will Cellular Slime Molds The life cycle of the protists called
require more supporting evidence to be widely accepted. cellular slime molds can prompt us to question what it means
to be an individual organism. The feeding stage of these organ-
isms consists of solitary cells that function individually, but
Amoebozoans when food is depleted, the cells form a slug-like aggregate that
The amoebozoan clade includes many species of amoebas functions as a unit (Figure 28.28). Unlike the feeding stage
that have lobe- or tube-shaped pseudopodia rather than the (plasmodium) of a plasmodial slime mold, these aggregated
threadlike pseudopodia found in rhizarians. Amoebozoans cells remain separated by their individual plasma membranes.
include tubulinids, slime molds, and entamoebas. Ultimately, the aggregated cells form an asexual fruiting body.
612 UNIT FIVE The Evolutionary History of Biological Diversity
. Figure 28.28 The life 1 In the feeding stage,
cycle of Dictyostelium, 9 In favorable solitary haploid amoebas 2 During sexual repro-
a cellular slime mold. conditions, amoebas engulf bacteria; these solitary duction, two haploid
emerge from the cells periodically divide by amoebas fuse and form
spore coats and feed. mitosis (asexual reproduction). a diploid zygote.

8 Spores
are released. FERTILIZATION
3 The zygote
7 Other Emerging becomes a giant
cells crawl amoeba cell by consuming
up the stalk Spores (n) haploid amoebas
and develop (n) (not shown). After
into spores. Zygote developing a
(2n) resistant wall, the
giant cell undergoes
600 om
Solitary amoebas meiosis followed by
(feeding stage) MEIOSIS several mitotic
(n) Amoebas divisions.
(n)
Fruiting
bodies
(n)
4 The wall ruptures,
releasing new
Aggregated haploid amoebas.
amoebas

Migrating
aggregate 5 When food is depleted,
hundreds of amoebas
congregate in response to a
chemical attractant and form
a slug-like aggregate
(see photo).

Key
6 The aggregate migrates for a 200 om
while and then stops. Some of the Haploid (n)
cells dry up after forming a stalk that
Diploid (2n)
supports a fruiting body.

VISUAL SKILLS Suppose cells were removed from the slug-like aggregate shown in the photo. Use
the information in the life cycle to infer whether these cells would be haploid or diploid. Explain.

Dictyostelium discoideum, a cellular slime mold commonly Entamoebas

found on forest floors, has become a model organism for Whereas most amoebozoans are free-living, those that belong
studying the evolution of multicellularity. One line of research to the genus Entamoeba are symbiotic parasites. They infect
has focused on the slime mold’s fruiting body stage. During all classes of vertebrate animals as well as some invertebrates.
this stage, the cells that form the stalk die as they dry out, Humans are host to at least six species of Entamoeba, but only
while the spore cells at the top survive and have the potential one, E. histolytica, is known to be pathogenic. E. histolytica
to reproduce (see Figure 28.28). Scientists have found that causes amoebic dysentery and is spread via contaminated
mutations in a single gene can turn individual Dictyostelium drinking water, food, or eating utensils. Responsible for up to
cells into “cheaters” that never become part of the stalk. 110,000 deaths worldwide every year, the disease is the third-
Because these mutants gain a strong reproductive advantage leading cause of death due to eukaryotic parasites, after malaria
over noncheaters, why don’t all Dictyostelium cells cheat? (see Figure 28.18) and schistosomiasis (see Figure 33.10).
Recent discoveries suggest an answer to this question.
Cheating cells lack a specific surface protein, and noncheat-
ing cells can recognize this difference. Noncheaters preferen- Opisthokonts
tially aggregate with other noncheaters, thus depriving cheat- Opisthokonts are an extremely diverse group of eukaryotes
ers of the chance to exploit them. Such a recognition system that includes animals, fungi, and several groups of protists.
may have been important in the evolution of other multicel- We will discuss the evolutionary history of fungi and animals
lular eukaryotes, such as animals and plants. in Chapters 31–34. Of the opisthokont protists, we will discuss
CHAPTER 28 Protists 613
the nucleariids in Chapter 31 because they are more closely c Figure 28.29
related to fungi than they are to other protists. Similarly, we A symbiotic
protist. This
will discuss choanoflagellates in Chapter 32, since they are
organism is a
more closely related to animals than they are to other protists. hypermastigote, a
The nucleariids and choanoflagellates illustrate why scientists member of a group
have abandoned the former kingdom Protista: A monophy- of parabasalids
that live in the gut
letic group that includes these single-celled eukaryotes would
of termites and
also have to include the multicellular animals and fungi that certain cockroaches
are closely related to them. and enable the
hosts to digest
CONCEPT CHECK 28.5 wood (SEM).

1. Contrast the pseudopodia of amoebozoans and forams.

10 om
2. In what sense is “fungus animal” a fitting description of a
slime mold? In what sense is it not fitting?
3. MAKE CONNECTIONS Alpha proteobacteria were used
as the outgroup to estimate the tree shown in Figure 28.26.
Explain why these bacteria were selected as the outgroup . Figure 28.30 Sudden oak death. Many dead oak trees are
rather than other bacterial or archaeal lineages. (See Figure visible in this Monterey County, California landscape. Infected trees
26.12 and associated text in Concept 26.3.) lose their ability to adjust to cycles of wet and dry weather.
For suggested answers, see Appendix A.

CONCEPT 28.6

Protists play key roles in

ecological communities
Most protists are aquatic, and they are found almost anywhere
there is water, including moist terrestrial habitats such as
damp soil and leaf litter. In oceans, ponds, and lakes, many
protists are bottom-dwellers that attach to rocks and other sub-
strates or creep through the sand and silt. As we’ve seen, other
protists are important constituents of plankton. We’ll focus species too. Massive fish kills have been attributed to Pfiesteria
here on two key roles that protists play in the varied habitats shumwayae (see Figure 28.17), a dinoflagellate parasite that
in which they live: that of symbiont and that of producer. attaches to its victims and eats their skin. Among species that
parasitize plants, the stramenopile Phytophthora ramorum
(an oomycete) has emerged as a major new forest pathogen.
Symbiotic Protists This species causes sudden oak death (SOD), a disease that
Many protists form symbiotic associations with other species. has killed millions of oaks and other trees in the United States
For example, photosynthetic dinoflagellates are food- and Great Britain (Figure 28.30; also see Concept 54.5).
providing symbiotic partners of the animals (coral polyps)
that build coral reefs. Coral reefs are highly diverse ecological
communities. That diversity ultimately depends on corals— Photosynthetic Protists
and on the mutualistic protists that nourish them. Corals Many protists are important producers, organisms that
support reef diversity by providing food to some species and use energy from light (or in some prokaryotes, inorganic
habitat to many others. chemicals) to convert CO2 to organic compounds. Producers
Another example is the wood-digesting protists that inhabit form the base of ecological food webs. In aquatic communi-
the gut of many termite species (Figure 28.29). Unaided, ter- ties, the main producers are photosynthetic protists and
mites cannot digest wood, and they rely on protistan or pro- prokaryotes (Figure 28.31). All other organisms in the com-
karyotic symbionts to do so. Termites cause over $3.5 billion munity are consumers that depend on producers for food,
in damage annually to wooden homes in the United States. either directly (by eating them) or indirectly (by eating an
Symbiotic protists also include parasites that have compro- organism that ate a producer). Scientists estimate that roughly
mised the economies of entire countries. Consider the malaria- 30% of the world’s photosynthesis is performed by diatoms,
causing protist Plasmodium: Income levels in countries hard dinoflagellates, multicellular algae, and other aquatic protists.
hit by malaria are 33% lower than in similar countries free Photosynthetic prokaryotes contribute another 20%, and
of the disease. Protists can have devastating effects on other plants are responsible for the remaining 50%.

614 UNIT FIVE The Evolutionary History of Biological Diversity

 A pressing question is how global warming will affect
photosynthetic protists and other producers. As shown in
Other Figure 28.32, the growth and biomass of photosynthetic pro-
consumers
tists and prokaryotes have declined in many ocean regions as
sea surface temperatures have increased. By what mechanism
Herbivorous do rising sea surface temperatures reduce the growth of marine
plankton Carnivorous
plankton producers? One hypothesis relates to the rise or upwelling
of cold, nutrient-rich waters from below. Many marine pro-
ducers rely on nutrients brought to the surface in this way.
However, rising sea surface temperatures can cause the for-
mation of a layer of light, warm water that acts as a barrier to
Prokaryotic Protistan nutrient upwelling—thus reducing the growth of marine pro-
producers producers ducers. If sustained, the changes shown in Figure 28.32 would
likely have far-reaching effects on marine ecosystems, fishery
yields, and the global carbon cycle (see Figure 55.14). Global
warming can also affect producers on land, but there the base
m Figure 28.31 Protists: key producers in aquatic
of food webs is occupied not by protists but by plants, which
communities. Arrows in this simplified food web lead from food
sources to the organisms that eat them. we will discuss in Chapters 29 and 30.

CONCEPT CHECK 28.6

Because producers form the foundation of food webs,
factors that affect producers can dramatically affect their 1. Justify the claim that photosynthetic protists are among the
entire community. In aquatic environments, photosynthetic biosphere’s most important organisms.
protists are often held in check by low concentrations of 2. Describe three symbioses that include protists.
nitrogen, phosphorus, or iron. Various human actions can 3. WHAT IF? High water temperatures and pollution can
increase the concentrations of these elements in aquatic cause corals to expel their dinoflagellate symbionts. How
might such “coral bleaching” affect corals and other species?
communities. For example, when fertilizer is applied to a
4. MAKE CONNECTIONS The bacterium Wolbachia is a symbiont
field, some of the fertilizer may be washed by rainfall into that lives in mosquito cells and spreads rapidly through mos-
a river that drains into a lake or ocean. When people add quito populations. Wolbachia can make mosquitoes resistant to
nutrients to aquatic communities in this or other ways, the infection by Plasmodium; researchers are seeking a strain that
confers resistance and does not harm mosquitoes. Compare
abundance of photosynthetic protists can increase spectacu-
evolutionary changes that could occur if malaria control is
larly. Such increases can have major ecological consequences, attempted using such a Wolbachia strain versus using insecti-
including the formation of large “dead zones” in marine cides to kill mosquitoes. (Review Figure 28.18 and Concept 23.4.)
ecosystems (see Figure 56.23). For suggested answers, see Appendix A.

. Figure 28.32 Effects of climate change on marine producers.

NP EP A Arctic (A)
NA
North Atlantic (NA)
NI EA Equatorial Atlantic (EA)
South Atlantic (SA)
Ocean region

North Indian (NI)

South Indian (SI)
North Pacific (NP)
Equatorial Pacific (EP)
South Pacific (SP)
SI SA Southern (S)
S SP
–0.02 –0.01 0.00 0.01 0.02
–2.50 –1.25 0.00 1.25 2.50
Chlorophyll change (mg/[m2 • yr])
Sea-surfacetemperature(SST)change(YC)
(a) Researchers studied 10 ocean regions, identified with letters (b) The concentration of chlorophyll, an index for the biomass and
on the map [see (b) for the corresponding names]. SSTs have growth of marine producers, has decreased over the same time
increased since 1950 in most areas of these regions. period in most ocean regions.

CHAPTER 28 Protists 615

28 Chapter Review Go to Mastering Biology for Assignments, the eText,
the Study Area, and Dynamic Study Modules.

• Current evidence indicates that eukaryotes originated by

SUMMARY OF KEY CONCEPTS endosymbiosis when an archaeal host (or a host closely re-
lated to the archaea) engulfed an alpha proteobacterium that
To review key terms, go to the Vocabulary Self-Quiz in the would evolve into an organelle found in all eukaryotes, the
Mastering Biology eText or Study Area, or go to [Link]/zkjz9t. mitochondrion.
• Plastids are thought to be descendants of cyanobacteria that
CONCEPT 28.1 were engulfed by early eukaryotic cells. The plastid-bearing
lineage eventually evolved into red algae and green algae.
Most eukaryotes are single-celled organisms (pp. 594–597) Other protist groups evolved from secondary endosymbiotic
• Domain Eukarya includes many groups of protists, along with events in which red algae or green algae were themselves
plants, animals, and fungi. Unlike prokaryotes, protists and other engulfed.
eukaryotes have a nucleus and other membrane-enclosed organ- • In one hypothesis, eukaryotes are grouped into four supergroups:
elles, as well as a cytoskeleton that enables them to have asym- Excavata, SAR, Archaeplastida, and Unikonta.
metric forms and to change shape as they feed, move, or grow.
• Protists are structurally and functionally diverse and have a wide ? Describe similarities and differences between protists and other
variety of life cycles. Most are unicellular. Protists include photo- eukaryotes.
autotrophs, heterotrophs, and mixotrophs.

Key Morphological
Key Concept/Eukaryote Supergroup Major Groups Characteristics Specific Examples

Diplomonads and Modified mitochondria Giardia,

CONCEPT 28.2
parabasalids Trichomonas
Excavates include protists with modified
Euglenozoans Spiral or crystalline rod Trypanosoma,
mitochondria and protists with unique inside flagella
Kinetoplastids Euglena
flagella (pp. 597–601)
Euglenids
? What evidence indicates that the excavates form a clade?

Stramenopiles Hairy and smooth flagella Phytophthora,

CONCEPT 28.3 Diatoms Laminaria
SAR is a highly diverse group of protists Oomycetes
defined by DNA similarities (pp. 601–609) Brown algae
Alveolates Membrane-enclosed sacs Pfiesteria,
? Although they are not photosynthetic, apicomplexan (alveoli) beneath plasma Plasmodium,
Dinoflagellates
parasites such as Plasmodium have modified plastids. Describe membrane Paramecium
a current hypothesis that explains this observation. Apicomplexans
Ciliates
Rhizarians Amoebas with threadlike Globigerina
Radiolarians pseudopodia
Forams
Cercozoans

CONCEPT 28.4 Red algae Phycoerythrin (photosyn- Porphyridium,

thetic pigment) Palmaria
Red algae and green algae are the closest
relatives of plants (pp. 609–611)
Green algae Plant-type chloroplasts Chlamydomonas,
? On what basis do systematists place plants in the same Ulva
supergroup (Archaeplastida) as red and green algae?

Plants (See Chapters 29 and 30.) Mosses, ferns,

conifers,
flowering
plants
Amoebozoans Amoebas with lobe- Amoeba, Dictyostelium
CONCEPT 28.5 Tubulinids shaped or tube-shaped
Unikonts include protists that are closely Slime molds pseudopodia
related to fungi and animals (pp. 611–614) Entamoebas Choanoflagellates,
Opisthokonts (Highly variable; see nucleariids,
? Describe a key feature for each of the main protist Chapters 31–34.) animals,
subgroups of Unikonta. fungi

616 UNIT FIVE The Evolutionary History of Biological Diversity

 CONCEPT 28.6 7. EVOLUTION CONNECTION • DRAW IT Medical researchers
seek to develop drugs that can kill or restrict the growth of
Protists play key roles in ecological human pathogens yet have few harmful effects on patients.
communities (pp. 614–615) These drugs often work by disrupting the metabolism of the
pathogen or by targeting its structural features.
• Protists form mutualistic and parasitic relationships that affect their Draw and label a phylogenetic tree that includes an
symbiotic partners and many other members of the community. ancestral prokaryote and the following groups of organisms:
• Photosynthetic protists are among the most important producers Excavata, SAR, Archaeplastida, Unikonta, and, within
in aquatic communities. Because they are at the base of the food Unikonta, amoebozoans, animals, choanoflagellates, fungi,
web, factors that affect photosynthetic protists affect many other and nucleariids. Based on this tree, hypothesize whether it
species in the community. would be most difficult to develop drugs to combat human
pathogens that are prokaryotes, protists, animals, or fungi.
? Describe several protists that are ecologically important.
(You do not need to consider the evolution of drug resistance
by the pathogen.)
TEST YOUR UNDERSTANDING
Levels 5-6: Evaluating/Creating
For more multiple-choice questions, go to the Practice Test in the 8. SCIENTIFIC INQUIRY Applying the “If . . . then” logic of
Mastering Biology eText or Study Area, or go to [Link]/GruWRg. deductive reasoning (see Concept 1.3), what are a few of the
predictions that arise from the hypothesis that plants evolved
Levels 1-2: Remembering/Understanding from green algae? Put another way, how could you test this
hypothesis?
1. Plastids that are surrounded by more than two membranes are
evidence of 9. WRITE ABOUT A THEME: INTERACTIONS Organisms
(A) evolution from mitochondria. interact with each other and the physical environment. In
(B) fusion of plastids. a short essay (100–150 words), explain how the response of
(C) origin of the plastids from archaea. diatom populations to a drop in nutrient availability can affect
(D) secondary endosymbiosis. both other organisms and aspects of the physical environment
(such as carbon dioxide concentrations).
2. Biologists think that endosymbiosis gave rise to mitochondria
before plastids partly because 10. SYNTHESIZE YOUR KNOWLEDGE
(A) the products of photosynthesis could not be metabolized
without mitochondrial enzymes.
(B) all eukaryotes have mitochondria (or their remnants),
whereas many eukaryotes do not have plastids.
(C) mitochondrial DNA is less similar to prokaryotic DNA than
is plastid DNA.
(D) without mitochondrial CO2 production, photosynthesis
could not occur.
3. Which group is correctly paired with its description?
(A) diatoms—important consumers in aquatic communities
(B) diplomonads—protists with modified mitochondria
(C) apicomplexans—producers with intricate life cycles
(D) red algae—acquired plastids by secondary endosymbiosis
4. According to the phylogeny presented in this chapter, which
protists are in the same eukaryotic supergroup as plants?

50 om
(A) green algae
(B) dinoflagellates
(C) red algae
(D) both A and C This micrograph shows a single-celled eukaryote, the ciliate
5. In a life cycle with alternation of generations, multicellular Didinium (left), about to engulf its Paramecium prey, which
haploid forms alternate with is also a ciliate. Identify the eukaryotic supergroup to which
(A) unicellular haploid forms. ciliates belong and describe the role of endosymbiosis in the
(B) unicellular diploid forms. evolutionary history of that supergroup. Are these ciliates
more closely related to all other protists than they are to plants,
(C) multicellular haploid forms.
fungi, or animals? Explain.
(D) multicellular diploid forms.
For selected answers, see Appendix A.
Levels 3-4: Applying/Analyzing
6. Based on the phylogenetic tree in Figure 28.5, which of the
following statements is correct?
(A) Excavata and SAR form a sister group.
(B) The most recent common ancestor of SAR is older than that
of Unikonta.
(C) The most basal (first to diverge) eukaryotic supergroup
cannot be determined.
(D) Excavata is the most basal eukaryotic supergroup.

CHAPTER 28 Protists 617