25

KEY CONCEPTS
The Origin and
Diversification of Eukaryotes
25.1 Eukaryotes Acquired Features
from Both Archaea and Bacteria
25.2 Major Lineages of Eukaryotes
Diversified in the Precambrian
25.3 Protists Reproduce Sexually
and Asexually
25.4 Protists Are Critical Components
of Many Ecosystems
© Don Paulson/PureStock/AGE Fotostock

A bloom of dinoflagellates of the genus Noctiluca

was responsible for this red tide in Puget Sound in
the U.S. state of Washington.

▶InvestigatingLIFE

Predicting Toxic Red Tides

In summer 2005, a devastating red tide crippled the shellfish future blooms, people in the area could be made aware of the
industry along the Atlantic coast of North America from Canada problem in advance and adjust the shellfish harvest (and their
to Massachusetts. This red tide was produced by a bloom of eating habits) accordingly.
dinoflagellates of the genus Alexandrium. These protists produce Biologists from WHOI monitored counts of dinoflagellates in
a powerful toxin that accumulates in clams, mussels, and oys- the water and in seafloor sediments. They also monitored river
ters. A person who eats a mollusk contaminated with the toxin runoff, water currents, water temperature and salinity, winds, and
can experience a syndrome known as paralytic shellfish poison- tides. An additional environmental factor was the “nor’easter”
ing. Many people were sickened by eating mollusks that were storms common along the New England coast. By correlating
harvested before the problem was diagnosed, and losses to the their measurements of these environmental factors with dino-
shellfish industry in 2005 were estimated at $50 million. flagellate counts, biologists produced a model that predicted
Several species of dinoflagellates produce toxic red tides growth of dinoflagellate populations.
in many parts of the world. Along the Gulf of Mexico, red tides In spring 2008, the WHOI team determined that all the factors
caused by dinoflagellates of the genus Karenia produce a neuro- were in place to produce another red tide like the one of 2005—if
toxin that affects the central nervous systems of fishes, which a nor’easter occurred to blow the dinoflagellates toward the coast.
become paralyzed and cannot respire effectively. Huge numbers A nor’easter did occur at just the wrong time, and another red tide
of dead fishes wash up on Gulf Coast beaches during a Karenia materialized in summer 2008, just as predicted. But this time,
red tide. In addition, wave action can produce aerosols of the people were warned. Shellfish harvesters adjusted their harvest,
Karenia toxin, and these aerosols often cause asthma-like symp- and many fewer people were harmed by eating toxic mollusks.
toms in humans on shore.
After the losses that resulted from the 2005 red tide, biolo-
gists at the Woods Hole Oceanographic Institution (WHOI) on
Cape Cod began to monitor and model dinoflagellate populations
QA
and
Red tides are harmful, but how are
dinoflagellates beneficial to marine ecosystems?
(Find the answer on page 578.)
off the New England coast. If biologists could accurately forecast

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 KEY CONCEPT 25.1 Eukaryotes Acquired Features from Both Archaea and Bacteria

• The origin of a nuclear envelope, which enclosed a genome

KEY CONCEPT Eukaryotes Acquired organized into chromosomes
Features from Both
25.1 Archaea and Bacteria
• The appearance of digestive vacuoles
• The acquisition of mitochondria and chloroplasts via
endosymbiosis
Learning Objectives
25.1.1 Explain the use of the term “protist” and describe FLEXIBLE CELL SURFACE We presume that ancient prokary-
the relationships of major protist groups to otic organisms, like most present-day prokaryotic cells, had firm
other eukaryotes. cell walls. The first step toward the eukaryotic condition was the
25.1.2 List the five key events that led to the evolution of loss of the cell wall by an archaean. This wall-less condition occurs
eukaryotes from a prokaryotic ancestor, and know in some present-day prokaryotes.
how these events allowed for the establishment of Consider the possibilities open to a flexible cell without a firm
unique cellular functions.
wall, starting with cell size. As a cell grows larger, its surface area-
25.1.3 Explain how major changes in Earth’s environment to-volume ratio decreases (see Figure 5.2). Unless the surface area
affected the evolution of the eukaryotic cell.
can be increased, the cell volume will reach an upper limit. If the
25.1.4 Classify the history of genes in a eukaryotic cell’s surface is flexible, however, it can fold inward and become
cell based on whether their origins are nuclear,
more elaborate, creating more surface area for gas and nutrient
mitochondrial, or chloroplastic.
exchange. With a surface flexible enough to allow infolding, the
cell can exchange materials with its environment rapidly enough
We easily recognize trees, mushrooms, and insects as plants, to sustain a larger volume and more rapid metabolism (see Fig-
fungi, and animals, respectively. But there is a dazzling assort- ure 25.1, steps 1–2). Furthermore, a flexible surface can pinch off
ment of other eukaryotic organisms—many microscopic, but bits of the environment, bringing them into the cell by endocyto-
some, like kelp, which grow very large—that do not fit into these sis. These infoldings of the cell surface, which also exist in some
three groups. Eukaryotes that are not plants, animals, or fungi modern prokaryotes, were important for the evolution of large
have traditionally been called protists. But phylogenetic analyses eukaryotic cells.
reveal that many of the groups we commonly refer to as protists
are not, in fact, closely related. Thus the term “protist” does not CHANGES IN CELL STRUCTURE AND FUNCTION Other early
describe a monophyletic group, but is a term of convenience for steps that were important for the evolution of the eukaryotic cell
“all the eukaryotes that are not plants, animals, or fungi.” involved increased compartmentalization and complexity of the
The unique characteristics of the eukaryotic cell lead scien- cell (see Figure 25.1, steps 3–7):
tists to conclude that the eukaryotes are monophyletic, and that
• The development of a more complex cytoskeleton
a single eukaryotic ancestor diversified into the many different
protist lineages as well as giving rise to the plants, fungi, and ani- • The formation of ribosome-studded internal membranes,
mals. As we saw in Key Concept 24.1, eukaryotes are a specialized some of which surrounded the DNA
group of archaea that acquired a cell nucleus. The mitochondria • The enclosure of the cell’s DNA in a nucleus
and chloroplasts of eukaryotes, however, are clearly derived from • The formation of a flagellum from microtubules of the
bacterial lineages (see Figure 24.1). cytoskeleton
Biologists have hypothesized that the origin of eukaryotes
• The evolution of digestive vacuoles
from a specialized archaeal ancestor was followed by the en-
dosymbioses with bacterial lineages that led to the origin of Until a few years ago, biologists thought that cytoskeletons were
mitochondria and chloroplasts. Some biologists prefer to view restricted to eukaryotes. Improved imaging technology and mo-
the origin of eukaryotes as the fusion of lineages from the two lecular analyses have now revealed homologs of many cytoskeletal
prokaryotic groups. This difference is largely a semantic one proteins in prokaryotes, so simple cytoskeletons evolved before the
that hinges on the subjective point at which we deem the eukary- origin of eukaryotes. The cytoskeleton of a eukaryote, however, is
otic lineage to have become definitively “eukaryotic.” In either much more developed and complex than that of a prokaryote. This
case, we can make some reasonable inferences about the events greater development of microfilaments and microtubules supports
that led to the evolution of a new cell type, bearing in mind that the eukaryotic cell and allows it to manage changes in shape, to
the environment underwent an enormous change—from low to distribute daughter chromosomes, and to move materials from
high availability of free atmospheric oxygen (O2)—during the one part of its larger cell to other parts. In addition, the presence
course of these events. of microtubules in the cytoskeleton allowed some cells to develop
the characteristic eukaryotic flagellum.
The modern eukaryotic cell arose in several steps The DNA of a prokaryotic cell is attached to a site on its cell
Several events were important in the origin of the modern eukary- membrane. If that region of the cell membrane were to fold into
otic cell (Figure 25.1): the cell, the first step would be taken toward the evolution of a
• The origin of a flexible cell surface
nucleus, a primary feature of the eukaryotic cell. The nuclear en-
velope appeared early in the eukaryotic lineage. The next step was
• The origin of a cytoskeleton probably phagocytosis—the ability to engulf and digest other cells.

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

Figure 25.1 Evolution of the Eukaryotic Cell The loss

Cell wall of a firm cell wall allowed the cell membrane to fold inward
1 The protective and create more surface area, which facilitated the evolution
cell wall was lost. DNA of larger cells. As cells grew larger, cytoskeletal complexity in-
creased, and the cell became increasingly compartmentalized.
Endosymbioses involving bacteria gave rise to mitochondria
and (in photosynthetic eukaryotes) to chloroplasts.
2 Infolding of the
cell membrane
Q: How could you use DNA sequencing and phyloge-
netic analysis to test the endosymbiotic origins
added surface area
hypothesis for mitochondria and chloroplasts?
without increasing
the cell’s volume.

ENDOSYMBIOSIS At the same time the processes we

3 Cytoskeleton
(microfilament have just outlined were taking place, cyanobacteria were
and microtubules) generating oxygen as a product of photosynthesis. The in-
increased in creasing concentrations of oxygen in the oceans, and even-
complexity.
tually in the atmosphere, had disastrous consequences for
most organisms of the time, which were unable to tolerate
4 Internal membranes
studded with
the newly oxidizing environment. But some prokaryotes
ribosomes formed. evolved strategies to use the increasing oxygen, and—fortu-
nately for us—so did some of the new phagocytic eukaryotes.
5 As regions of the At about this time, endosymbiosis began to play a role
infolded cell in eukaryotic evolution (see Figure 25.1, steps 8–9). The
membrane enclosed theory of endosymbiosis proposes that certain organelles
the cell’s DNA, a
are the descendants of prokaryotes engulfed, but not di-
precursor of a
nucleus formed. gested, by ancient eukaryotic cells. One crucial event in the
6 Microtubules from
the cytoskeleton
history of eukaryotes was the incorporation of a proteobac-
7 Early digestive formed the terium that evolved into the mitochondrion. Initially the
vacuoles evolved eukaryotic flagellum, new organelle’s primary function was probably to detoxify
into lysosomes using enabling propulsion. O2 by reducing it to water. Later this reduction became
enzymes from the early coupled with the formation of ATP in cellular respiration.
endoplasmic reticulum.
After this step, the essential eukaryotic cell was complete.

8 Mitochondria formed
Connect the Concepts You may wish to review the
through endosymbiosis reactions of cellular respiration in Key Concept 5.3.
with a proteobacterium.
Photosynthetic eukaryotes are the result of yet another
endosymbiotic step: the incorporation of a prokaryote relat-
9 Endosymbiosis with ed to today’s cyanobacteria, which became the chloroplast.
cyanobacteria led to
the development of Chloroplasts have been transferred among
chloroplasts.
eukaryotes several times
Eukaryotes in several different groups possess chloroplasts,
and groups with chloroplasts appear in several distantly
related eukaryotic clades. Some of these groups differ in
the photosynthetic pigments their chloroplasts contain.
And not all chloroplasts are limited to a pair of surround-
Flagellum ing membranes—in some microbial eukaryotes, chloro-
Mitochondrion plasts are surrounded by three or more membranes. We
Chloroplast Nucleus now view these observations as evidence of a remarkable
series of endosymbioses. This conclusion is supported by
extensive evidence from electron microscopy and nucleic
acid sequence comparisons.
All chloroplasts trace their ancestry back to the engulf-
ment of one cyanobacterium by a larger eukaryotic cell.
This event, the step that first gave rise to the photosynthetic
eukaryotes, is known as primary endosymbiosis (Figure
25.2A). The cyanobacterium, a Gram-negative bacterium,

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 KEY CONCEPT 25.1 Eukaryotes Acquired Features from Both Archaea and Bacteria

(A) Primary endosymbiosis following primary endosymbiosis (as we will see in Chapter 26).
Eukaryote Cyanobacterium Primary endosymbiosis also gave rise to the chloroplasts of the
Peptidoglycan red algae, green algae, and land plants. The red algal chloroplast
retains certain pigments of the original cyanobacterial endosym-
Cyanobacterium biont that are absent in green algal chloroplasts.
outer membrane Over evolutionary Almost all remaining photosynthetic eukaryotes are the re-
Cyanobacterium history, one of
these three
sult of additional rounds of endosymbiosis. For example, the
inner membrane
membranes photosynthetic euglenids derived their chloroplasts from sec-
Host cell was lost. ondary endosymbiosis (Figure 25.2B). Their ancestor took up
membrane
Host cell a unicellular green alga, retaining its chloroplast and eventually
nucleus losing the rest of the constituents of the alga. This history ex-
Chloroplast plains why the photosynthetic euglenids have the same photo-
synthetic pigments as the green algae and land plants. It also
Peptidoglycan has
accounts for the third membrane of the euglenoid chloroplast,
been lost except in which is derived from the euglenid’s cell membrane (as a result
glaucophytes. of endocytosis). An additional round of endosymbiosis—tertiary
endosymbiosis—occurred when a dinoflagellate apparently lost
its chloroplast and took up another protist that had acquired its
chloroplast through secondary endosymbiosis.
KEY CONCEPT
(B) Secondary endosymbiosis
Chloroplast- 25.1 Recap and Assess
Host eukaryotic cell
containing
eukaryotic cell The modern eukaryotic cell probably arose from an ancestral
prokaryotic archaean in several steps, including the origin of a
flexible cell surface and the enclosure of the genetic material in
a nucleus. Later, endosymbioses of proteobacteria and cyano-
bacteria led to the origins of mitochondria and chloroplasts,
respectively.
1. Why was the development of a flexible cell surface a key
event for eukaryotic evolution?
2. Explain how increased availability of atmospheric oxygen (O2)
Host membrane (from could have influenced the evolution of the eukaryotic cell.
endocytosis) encloses
the engulfed cell.
Background for Questions 3–4: Ribosomal RNA (rRNA) genes
are present in the nuclear genome of eukaryotes. There are
A trace of the engulfed also rRNA genes in the genomes of mitochondria and chloro-
cell’s nucleus is retained plasts. Therefore photosynthetic eukaryotes have three differ-
in some groups. ent sets of rRNA genes, which encode the structural RNA of
three separate sets of ribosomes. The gene tree below shows
the evolutionary relationships among rRNA gene sequences
The inner of these two isolated from the nuclear genomes of humans, yeast, and corn;
membranes has been
from a prokaryotic archaean (Halobacterium), a proteobacterium
lost in euglenids
(E. coli), and a cyanobacterium (Chlorobium); and from the mito-
and dinoflagellates.
chondrial and chloroplast genomes of corn.

Figure 25.2 Endosymbiotic Events in the Evolution of Chloroplasts Halobacterium rRNA

(A) A single instance of primary endosymbiosis ultimately gave rise to all
of today’s chloroplasts. (B) Secondary endosymbiosis—the uptake and Human nuclear rRNA
retention of a chloroplast-containing cell by another eukaryotic cell—took
place several times, independently. Yeast nuclear rRNA
View in Achieve
Corn nuclear rRNA
Animation 25.1 Family Tree of Chloroplasts
E. coli rRNA
had both an inner and an outer membrane (see Figure 24.2B). Thus
Corn mitochondrial rRNA
the original chloroplasts had two surrounding membranes: the in-
ner and outer membranes of the cyanobacterium. Remnants of the Chlorobium rRNA
peptidoglycan-containing cell wall of the bacterium are present in
the form of a bit of peptidoglycan between the chloroplast mem- Corn chloroplast rRNA
branes of glaucophytes, the first eukaryotic group to branch off

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

3. Why are the three rRNA genes of corn not one another’s opiles, rhizarians, excavates, plants, amoebozoans, fungi, and
closest relatives?
animals (Figure 25.3). Plants, fungi, and animals each have close
4. How would you explain the closer relationship of the protist relatives (such as the choanoflagellate relatives of animals),
mitochondrial rRNA gene of corn to the rRNA gene of E. coli
than to the nuclear rRNA genes of other eukaryotes? Can you which we will discuss along with those major multicellular eukary-
explain the relationship of the rRNA gene from the chloroplast otic groups in Chapters 26–31.
of corn to the rRNA gene of the cyanobacterium? Each of the five major groups of protist eukaryotes covered
5. Explain why the term “protists” does not refer to a formal in this chapter consists of organisms with enormously diverse
taxonomic group. body forms and nutritional lifestyles. Some protists are motile,
whereas others do not move. Some protists are photosynthetic,
The features that eukaryotes gained from prokaryotic archaea and whereas others are heterotrophic. Most protists are unicellular,
bacteria have allowed them to exploit many different environ- but some are multicellular. Most protists are microscopic, but a
ments. This led to the evolution of great diversity among eukary- few are huge (giant kelps, for example, can grow to half the length
otes, beginning with a radiation that started in the Precambrian. of a football field). We refer to the unicellular species of protists
as microbial eukaryotes, but keep in mind that there are large,
multicellular protists as well.
KEY CONCEPT Major Lineages of Eukaryotes Multicellularity has arisen dozens of times across the evolution-

25.2 Diversified in the Precambrian

ary history of eukaryotes. Four of the origins of multicellularity
resulted in large organisms that are familiar to most people: plants,
animals, fungi, and brown algae (the last are a group of strameno-
Learning Objectives
piles). In addition, there are dozens of smaller and less familiar
25.2.1 Distinguish among the major groups of microbial groups among the eukaryotes that include multicellular species.
eukaryotes (alveolates, stramenopiles, rhizarians, Recent experimental studies have shown that artificial selection
excavates, and amoebozoans).
for multicellularity can produce repeated, convergent evolution of
25.2.2 Justify the position that it is relatively easy for multicellular forms over just a few months in some normally unicel-
multicellularity to evolve.
lular eukaryotic species. In addition, many unicellular species retain
25.2.3 Elaborate on why protists are important to our individual identities but nonetheless associate in large multicellu-
medical and economic concerns.
lar colonies. There is a near-continuum between fully integrated,
multicellular organisms on the one hand and loosely integrated
Most eukaryotes can be placed in one of eight major clades that multicellular colonies of cells on the other. Biologists do not always
began to diversify about 1.5 billion years ago: alveolates, stramen- agree on where to draw the line between the two.

Alveolates

Stramenopiles

Rhizarians

Excavates

Plantae Chs. 26 and 27

Amoebozoans

Fungi Ch. 28

Choanoflagellates
Ophisthokonts Chs.
29–31
Animals

Precambrian Paleozoic Mesozoic Cenozoic

to 1.5 1.4 1.3 1.2 1.1 1.0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0
4.5 bya Time (bya)

Figure 25.3 Precambrian Divergence of Major Eukaryotic Groups major lineages between 1.5 and 1.4 billion years ago (bya) makes recon-
A phylogenetic tree shows a current hypothesis and estimated time line struction of their precise relationships difficult. The major multicellular
for the origin of the major groups of eukaryotes. The rapid divergence of groups (tinted boxes) will be covered in subsequent chapters.

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 KEY CONCEPT 25.2 Major Lineages of Eukaryotes Diversified in the Precambrian

Biologists used to classify protists largely on the basis of their Amphidiniopsis kofoidii
life histories and reproductive features. In recent years, however,
electron microscopy and gene sequencing have revealed many
new patterns of evolutionary relatedness among these groups.
Analyses of slowly evolving gene sequences are making it pos-
sible to explore evolutionary relationships among eukaryotes
in ever greater detail and with greater confidence. Nonetheless,
some substantial areas of uncertainty remain, and lateral gene
transfer may complicate efforts to reconstruct the evolutionary
history of protists (as was also true for prokaryotes; see Key

© Biophoto Associates/Science Source

Concept 24.1). Today we recognize great diversity among the
many distantly related protist clades.

Alveolates have sacs Dinoflagellates

Alveolates
under their cell Apicomplexans
membranes
Ciliates
Alveolates are so named be-
Equatorial groove Flagellum Longitudinal groove 20 µm
cause they possess sacs, called Stramenopiles

alveoli, just beneath their cell Rhizarians Figure 25.4 A Dinoflagellate The presence of two flagella is
membranes, which may play a characteristic of many dinoflagellates, although these appendages
role in supporting the cell surface. All alveolates are unicellular, are contained within deep grooves and thus are seldom visible. One
and most are photosynthetic, but they are diverse in body form. flagellum lies within the equatorial groove and provides forward thrust
The alveolate groups we consider in detail here are the dinoflagel-
Alveolates and spin to the organism. The second flagellum originates in the
lates, apicomplexans, and ciliates. longitudinal groove and acts like the rudder of a boat.
Diatoms
Stramenopiles

Media Clip 25.1 A Dinoflagellate Shows Off Its Flagellum

DINOFLAGELLATES Most dinoflagellates are marineBrown andalgae
pho- Life12e.com/mc25.1
tosynthetic; they are important primary producers ofOomycetes organic
matter in the oceans. Although fewer species of dinoflagellates
live in fresh water, individuals can be abundant in freshwater Rhizarians
the apical complex enables Plasmodium, the causative agent of
environments. The dinoflagellates are of great ecological, evo- malaria, to enter its target cells in the human body after trans-
lutionary, and morphological interest. A distinctive mixture of mission by a mosquito.
photosynthetic and accessory pigments gives their chloroplastsAlveolates
a Like many obligate parasites, apicomplexans have elaborate
golden brown color. Some dinoflagellate species cause red tides, life cycles featuring asexual and sexual reproduction through a
as discussed at the start of this chapter. Other species are pho-
Stramenopiles series of very dissimilar life stages (see Figure 25.20). In many
tosynthetic endosymbionts that live within the cellsCercozoansof other species, these life stages are associated with two different types
organisms, including invertebrate animals (such as corals; see of host organisms, as is the case with Plasmodium. Another
Rhizarians

Foraminiferans
Investigating Life: Can Corals Reacquire Dinoflagellate Endo- apicomplexan, Toxoplasma, alternates between cats and rats to
symbionts Lost to Bleaching?) and other marine protists (see
Radiolarians complete its life cycle. A rat infected with Toxoplasma loses its
Figure 25.12A). Still others are nonphotosynthetic and live as fear of cats, which makes it more likely to be eaten by, and thus
parasites within other marine organisms. transfer the parasite to, a cat.
Dinoflagellates have a distinctive appearance. They generally
Excavates
have two flagella, one in an equatorial groove around the cell, the
Diplomonads CILIATES The ciliates are named for their numerous hairlike
other starting near the same point as the first andParabasalids
passing down cilia, which are shorter than, but otherwise identical to, eukaryotic
a longitudinal groove before extending into the surrounding me- flagella. The ciliates are much more complex in body form than
Heteroloboseans
dium (Figure 25.4). Some dinoflagellates can take on different are most other unicellular eukaryotes (Figure 25.5). Their defini-
forms, including amoeboid ones, depending on environmental
Euglenids tive characteristic is the possession of two types of nuclei (whose
conditions. It has been claimed that the dinoflagellate Pfiesteria roles we describe in Key Concept 25.3 when we discuss protist
Kinetoplastids
piscicida can occur in at least two dozen distinct forms, although reproduction). Almost all ciliates are heterotrophic, although a
this claim is highly controversial. In any case, this remarkable few contain photosynthetic endosymbionts.
dinoflagellate, when present in large enough numbers, is harmful Paramecium, a frequently studied ciliate genus, exemplifies
Amoebozoans
to fishes and can both stun and feed on them. Loboseans the complex structure and behavior of ciliates (Figure 25.6).
The slipper-shaped cell is covered by an elaborate pellicle, a
Plasmodial slime molds
APICOMPLEXANS The exclusively parasitic apicomplexans structure composed principally of an outer membrane and an
derive their name from the apical complex, aCellular
massslime
of organelles
molds inner layer of closely packed, membrane-enclosed sacs (the al-
contained in the apical end (the tip) of the cell. These organelles veoli) that surround the bases of the cilia. Defensive organelles
help the apicomplexan invade its host’s tissues. For example, called trichocysts are also present in the pellicle. In response to

Life 12e
Oxford University Press
Dragonfly Media Group
Life12e_25.04.ai Date 06-17-19

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

(A) Paramecium sp. (B) Didinium nasutum (C) Trichodina sp.

Oral surface

Cilia

© Aaron Bell/Visuals Unlimited, Inc.

© Eye of Science/Science Source

© SPL/Science Source

Cilia 10 µm Bands of cilia 7 µm 15 µm

Figure 25.5 Diversity among the Ciliates (A) A free-swimming nasutum feeds on other ciliates, including Paramecium. Its cilia occur in
organism, this Paramecium belongs to a ciliate group whose members two separate bands. (C) The parasite Trichodina creates suction with its
have many cilia of uniform length. (B) The barrel-shaped Didinium ring of cilia that help attach it to the surface of a fish host.

a threat, a microscopic explosion expels the trichocysts in a few Organisms living in fresh water are hypertonic to their envi-
milliseconds, and they emerge as sharp darts, driven forward at ronment. Many freshwater protists, including Paramecium, ad-
the tip of a long, expanding filament. dress this problem by means of specialized contractile vacuoles
The cilia provide Paramecium with a form of locomotion that that excrete the excess water the organisms constantly take in by
is generally more precise than locomotion by flagella. A Parame- osmosis. The excess water collects in the contractile vacuoles,
cium can coordinate the beating of its cilia to propel itself either which then contract and expel the water from the cell.
forward or backward in a spiraling manner. It can also back off Paramecium and many other protists engulf solid food by en-
swiftly when it encounters a barrier or a negative stimulus. The docytosis, forming a digestive vacuole within which the food is
coordination of ciliary beating is probably the result of a dif- digested (Figure 25.7). Smaller vesicles containing digested food
ferential distribution of ion channels in the cell membrane near pinch away from the digestive vacuole and enter the cytosol. These
the two ends of the cell. tiny vesicles provide a large surface area across which the products
of digestion can be absorbed by the rest of the cell.
View in Achieve
Animation 25.2 Digestive Vacuoles
Micronuclei function in The macronucleus controls
genetic recombination. the cell’s activities. Stramenopiles Alveolates

typically have two Diatoms

Stramenopiles
Contractile unequal flagella, one
vacuole with hairs Brown algae

A morphological synapomor- Oomycetes

Alveoli phy of most stramenopiles
Rhizarians
is the possession of rows of
Cilia tubular hairs on the longer of their two flagella. Some stramenopiles
Digestive lack flagella, but they are descended from ancestors that possessed
vacuole
flagella. The stramenopiles include the diatoms and the brown al-
Oral groove
gae, which are photosynthetic, and the oomycetes, which are not.
Anal pore Trichocyst
DIATOMS All of the diatoms are unicellular, although some
Fibrils
Pellicle species associate in filaments. Many have sufficient carotenoids in
Alveolus their chloroplasts to give them a yellow or brownish color. All of
Cilium them synthesize carbohydrates and oils as photosynthetic storage
products. Diatoms lack flagella except in male gametes.
Architectural magnificence on a microscopic scale is the hall-
Figure 25.6 Anatomy of Paramecium Paramecium, with its many mark of the diatoms. Almost all diatoms deposit silica (hydrated
specialized organelles, exemplifies the complex body form of ciliates. silicon dioxide) in their cell walls. The cell wall of a diatom is
View in Achieve constructed in two pieces, with the top overlapping the bottom
Activity 25.1 Anatomy of Paramecium like the top of a petri dish. The silica-impregnated walls have

25_Life12e_Ch 25.indd 7 12/3/19 4:10 PM

 KEY CONCEPT 25.2 Major Lineages of Eukaryotes Diversified in the Precambrian

Experiment
Figure 25.7 The Role of Vacuoles in Ciliate Digestion
Original Paper: S. O. Mast. 1947. The food-vacuole in Parame-
cium. Biol Bull 92: 31–72.
An acidic environment is known to aid digestion in many multicel-

HYPOTHESIS▸ The digestive vacuoles of Paramecium produce

lular organisms. Do ciliates also use acid to obtain nutrients?

© James Solliday/Biological Photo Service

an acidic environment that allows the organism to digest food

METHOD▸
particles.

1. Feed Paramecium yeast cells stained with Congo red, a dye

that is red at neutral or basic pH but turns green at acidic pH.
2. Under a light microscope, observe the formation and degrada-
tion of digestive vacuoles within the Paramecium. Note time

RESULTS▸
and sequence of color (i.e., acid level) changes.

Diatoms display either radial …or bilateral 25 µm

(circular) symmetry… (left–right) symmetry.
1 A digestive vacuole forms
around yeast cells.
Figure 25.8 Diatom Diversity This bright-field micrograph illus-
trates the variety of species-specific forms found among the diatoms.
2 The change in color Media Clip 25.2 Diatoms in Action
shows that the interior
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vacuole has become
Stained acidic.
yeast cells
fuse. The resulting zygote then grows substantially in size before
a new cell wall is laid down.
Diatoms are found in all the oceans and are frequently present
Oral 3 As products of in great numbers. They are major photosynthetic producers in
groove digestion move into
the cytosol, the pH
coastal waters (see Key Concept 25.4) and are among the dominant
increases in the organisms in the dense “blooms” of phytoplankton that occasion-
vacuole (the dye ally appear in the open ocean. Diatoms are also common in fresh
becomes red again). water and even occur on the wet surfaces of terrestrial mosses.

BROWN ALGAE The brown algae obtain their namesake color

4 Red-stained from the carotenoid fucoxanthin, which is abundant in their chlo-
(basic) waste roplasts. The combination of this yellow-orange pigment with the
material is
expelled.
green of chlorophylls a and c yields a brownish tinge. All brown

CONCLUSION▸ Some ciliates acidify digestive vacuoles to

algae are multicellular, and some are extremely large. Giant kelps
such as those of the genus Macrocystis may be up to 60 meters long.
assist in the breakdown of food. Media Clip 25.3 A Kelp Forest
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The brown algae are almost exclusively marine. They are
composed either of branched filaments (Figure 25.9A) or of
intricate patterns unique to each species (Figure 25.8). Despite leaflike growths (Figure 25.9B). Some float in the open ocean;
their remarkable morphological diversity, all diatoms are sym- the most famous example is the genus Sargassum, which forms
metrical—either bilaterally (with “right” and “left” halves) or radi- dense mats in the Sargasso Sea in the mid-Atlantic. Most brown
ally (with the type of symmetry possessed by a circle). algae, however, attach themselves to rocks near the shore. A few
Diatoms reproduce both sexually and asexually. Asexual re- thrive only where they are regularly exposed to heavy surf. All
production by binary fission is somewhat constrained by the stiff of the attached forms develop a specialized structure, called a
cell wall. Both the top and bottom of the “petri dish” become holdfast, that literally glues them to the rocks. The “glue” of the
tops of new “dishes” without changing appreciably in size. As a holdfast is alginic acid, a gummy polymer found in the walls
result, the new cell made from the former bottom is smaller than of many brown algal cells. In addition to its function in hold-
the parent cell. If this process continued indefinitely, one cell line fasts, alginic acid cements algal cells and filaments together. It
would simply vanish, but sexual reproduction largely solves this is harvested and used by humans as an emulsifier in ice cream,
potential problem. Gametes are formed, shed their cell walls, and cosmetics, and other products.

25_Life12e_Ch 25.indd 8 12/3/19 4:10 PM

 Dinoflagellates

Alveolates
Apicomplexans

Ciliates

CHAPTER 25 The Origin and Diversification of Eukaryotes Stramenopiles

Rhizarians

(A) Filamentous growth pattern (B) Leaf-like growth pattern Some other oomycetes, such as the downy mildews, are ter-
restrial. Although most of the terrestrial oomycetes are harmless
Alveolates
or helpful decomposers of dead matter, a few are plant parasites
Diatoms
that attack crops such as avocados, grapes, and potatoes.

Stramenopiles
Oomycetes were once classified as fungi. However, we now
Brown algae
know that their similarity to fungi is only superficial, and that the
Oomycetes
oomycetes are more distantly related to the fungi than are many
other eukaryotic groups, including humans (see Figure 25.3). For
Rhizarians
example, the cell walls of oomycetes are typically made of cel-
lulose, whereas those of fungi are made of chitin.

© Carl W. May/Biological Photo Service

Rhizarians typically Alveolates
© Marevision/AGE Fotostock

have long, thin Stramenopiles

pseudopods
Cercozoans
The three primary groups

Rhizarians
of rhizarians—cercozoans, Foraminiferans

foraminiferans, and radio- Radiolarians

Himanthalia elongata Postelsia palmaeformis Holdfasts
larians—are unicellular and
Figure 25.9 Brown Algae (A) This seaweed illustrates the fila- mostly aquatic. The rhizarians have contributed their shells to
mentous growth form of the brown algae. (B) Sea palms exemplify the ocean sediments, some of which have become terrestrial features
leaflike growth form of brown algae. Sea palms and many other brown over the course of geological history.
Excavates
Diplomonads
algal species are “glued” to the rocks by tough, branched structures
called holdfasts that can withstand the pounding of the surf. Parabasalids
CERCOZOANS The cercozoans are a diverse group with many
forms and habitats. Some are aquatic; others live in soil. One group
Heteroloboseans

OOMYCETES The oomycetes are the water molds and their of cercozoans possesses chloroplasts derived fromEuglenids
a green alga by
terrestrial relatives. Water molds are filamentous and stationary. secondary endosymbiosis, and those chloroplasts contain a trace
They are absorptive heterotrophs—that is, they secrete enzymes of the alga’s nucleus. Kinetoplastids

that digest large food molecules into smaller molecules that they
can absorb. They are all aquatic and saprobic—meaning they feed FORAMINIFERANS Some foraminiferans secrete external shells
on dead organic matter. If you have seen a whitish, cottony mold of calcium carbonate (Figure 25.11). These shells have accumu-
Amoebozoans
Loboseans
growing on dead fishes or dead insects in water, it was probably lated over time to produce much of the world’s limestone. Some
a water mold of the common genus Saprolegnia (Figure 25.10). foraminiferans live as plankton; others live on the
Plasmodial seafloor.
slime molds Living
Cellular slime molds

Saprolegnia sp.

© Robert Brons/Biological Photo Service

David McIntyre

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3 mm
Figure 25.11 Building Blocks of Limestone Some foraminiferans
secrete calcium carbonate to form shells. The shells of different
Figure 25.10 An Oomycete The filaments of a water mold radiate species have distinctive shapes. Over millions of years, the shells of
from the carcass of a beetle. foraminiferans have accumulated to form limestone deposits.

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 KEY CONCEPT 25.2 Major Lineages of Eukaryotes Diversified in the Precambrian

(A) Amphilonche heteracantha (B) Thyrsocyrtis sp. mitochondria that were lost or reduced over the course of
evolution. The existence of these organisms today shows
that eukaryotic life is possible without mitochondria, at
least among parasitic species.

DIPLOMONADS AND PARABASALIDS The diplomo-

© Steve Gschmeissner/Science Source

nads and the parabasalids are unicellular and lack mito-
chondria (although they have reduced organelles that are
derived from mitochondria). The parasitic Giardia lam-
blia, a diplomonad, causes the intestinal disease giardiasis.

© Larry Jon Friesen

Giardia infections may result from contact with contami-
nated water. In the United States, such infections are com-
mon among hikers and campers using spring or stream
50 µm 100 µm water in recreational areas, as well as among children kept
in close quarters (as in day-care centers). The tiny Giardia
Figure 25.12 Radiolarians Exhibit Distinctive Pseudopods and Radial
Dinoflagellates
contain two nuclei bounded by nuclear envelopes and have
Symmetry (A) The radiolarians are distinguished by their thin, stiff pseudo-

Alveolates
pods and by their radial symmetry. The pigmentation seen Apicomplexans
at the center of a cytoskeleton and multiple flagella (Figure 25.13A).
this radiolarian’s glassy endoskeleton is imparted by endosymbiotic dinofla- In addition to flagella and a cytoskeleton, the parabasa-
Ciliates
gellates. (B) The endoskeleton secreted by a radiolarian. lids have undulating membranes that also contribute to the
Stramenopiles cell’s locomotion. Trichomonas vaginalis (Figure 25.13B)
Rhizarians
is a parabasalid responsible for a sexually transmitted dis-
foraminiferans have been found 10,896 meters down in the west- ease in humans. Infection of the male urethra, where it may occur
ern Pacific’s Challenger Deep—the deepest point in the world’s without symptoms, is less common than infection of the vagina.
oceans. At that depth, however, they cannot secrete normal shells
because the surrounding water is too poor in calcium carbonate. Alveolates
(A) Giardia muris
In living planktonic foraminiferans, long, threadlike, branched
Diatoms
Stramenopiles

pseudopods extend through numerous microscopic apertures in

the shell and interconnect to create a sticky, reticulated net, which
Brown algae

the foraminiferans use to catch smaller plankton. In some forami- Oomycetes

niferan species, the pseudopods provide locomotion.
Rhizarians

Courtesy of Dr. Stan Erlandsen/CDC

RADIOLARIANS Radiolarians are recognizable by their thin,
stiff pseudopods, which are reinforced by microtubules (Figure
25.12A). These pseudopods greatly increase the surface area ofAlveolates
the
cell, and they help the cell stay afloat in its marine environment.
Radiolarians also are immediately recognizable by their dis-
Stramenopiles

tinctive radial symmetry. Almost all radiolarian species secreteCercozoans

glassy endoskeletons (internal skeletons). The skeletons of the
Rhizarians

Foraminiferans 2 µm
different species are as varied as snowflakes, and many have (B) Trichomonas vaginalis
elaborate geometric designs (Figure 25.12B). A few radiolariansRadiolarians
are among the largest of the unicellular eukaryotes, measuring
several millimeters across.
Excavates
Excavates began to diversify Diplomonads

about 1.5 billion years ago

© Eye of Science/Science Source

Parabasalids
The excavates include several di-
Heteroloboseans
verse groups that began to split from
one another soon after the origin of Euglenids

eukaryotes. Several groups of exca-

Kinetoplastids
vates lack mitochondria, an absence
that once led to the view that these groups might represent early-
diverging eukaryotes that diversified before the evolution of mito- 2 µm
Amoebozoans
chondria. However, the discovery of genes in the nucleusLoboseansthat are Figure 25.13 Some Excavate Groups Lack Mitochondria
normally associated with mitochondria suggests that the absence (A) Giardia, a diplomonad, has flagella and two nuclei. (B) Trichomonas,
Plasmodial slime molds
of mitochondria is a derived condition in these organisms. In oth- a parabasalid, has flagella and undulating membranes. Neither of
er words, ancestors of these excavate groups probablyCellularpossessed
slime molds these organisms possess mitochondria.

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

Photosynthetic chloroplasts are prominent photosynthetic pigment and begin to feed exclusively on dissolved
features in a typical Euglena cell. organic material in the water around them. A “bleached” Euglena
resynthesizes its photosynthetic pigment when it is returned to the
light and becomes autotrophic again. But Euglena cells treated with
Flagella Nucleus certain antibiotics or mutagens lose their photosynthetic pigment
completely; neither they nor their descendants are ever autotrophs
Pigment again. However, those descendants function well as heterotrophs.
shield
The kinetoplastids are unicellular parasites with two flagella
and a single, large mitochondrion. The mitochondrion contains
a kinetoplast, a unique structure housing multiple circular DNA
molecules and associated proteins. Some of these DNA molecules
encode “guide proteins” that edit mRNA within the mitochondrion.
Contractile Stored polysaccharides The kinetoplastids include several medically important species
Photoreceptor vacuole from photosynthesis of pathogenic trypanosomes (Table 25.1). Some of these organ-
isms are able to change their cell surface recognition molecules
Figure 25.14 A Photosynthetic Euglenid In the Euglena species
illustrated in this drawing, the second flagellum is rudimentary. Note that
frequently, allowing them to evade our best attempts to kill them
the primary flagellum originates at the anterior of the organism and trails and thus eradicate the diseases they cause.
toward its posterior. Amoebozoans
Amoebozoans use Loboseans
lobe-shaped pseudopods
HETEROLOBOSEANS The amoeboid body form appears in for locomotion Plasmodial slime molds

several protist groups that are only distantly related to one an- Amoebozoans appear to have di- Cellular slime molds

other. The body forms of heteroloboseans, for example, resemble verged from other eukaryotes about
those of loboseans, an amoebozoan group that is not at all closely 1.5 billion years ago (see Figure 25.3). It is not yet clear whether
related to heteroloboseans (see the next section). Amoebas of the they are more closely related to opisthokonts (which include fungi
free-living heterolobosean genus Naegleria, some of which can and animals) or to other major groups of eukaryotes.
enter the human body and cause a fatal disease of the nervous The lobe-shaped pseudopods of amoebozoans (Figure 25.15)
system, usually have a two-stage life cycle, in which one stage has are a hallmark of the amoeboid body form. Amoebozoan pseu-
amoeboid cells and the other flagellated cells. dopods differ in form and function from the slender pseudopods
of rhizarians. We consider three amoebozoan groups here: the
EUGLENIDS AND KINETOPLASTIDS The euglenids and kineto- loboseans and two groups known as slime molds.
plastids together constitute a clade of unicellular excavates with
flagella. Their mitochondria contain distinctive disc-shaped cris- LOBOSEANS Loboseans are small amoebozoans that feed on
tae, and their flagella contain a crystalline rod not found in other other small organisms and particles of organic matter by phago-
organisms. They reproduce primarily asexually by binary fission. cytosis, engulfing them with pseudopods. Many loboseans are
The flagella of euglenids arise from a pocket at the anterior adapted for life on the bottoms of lakes, ponds, and other bodies
end of the cell. Spiraling strips of proteins under the cell mem- of water. Their creeping locomotion and their manner of engulf-
brane control the cell’s shape. Some euglenids are photosyn- ing food particles fit them for life close to a relatively rich supply
thetic. Figure 25.14 depicts a typical cell of the genus Euglena. of sedentary organisms or organic particles. Most loboseans ex-
Like most other euglenids, this common freshwater organism has ist as predators, parasites, or scavengers. Members of one group
a complex cell structure. It propels itself through the water with of loboseans, the testate amoebas, live inside shells. Some of
the longer of its two flagella, which
may also serve as an anchor to hold
the organism in place. The second TABLE 25.1 | Three Pathogenic Trypanosomes
flagellum is often rudimentary. Trypanosoma brucei Trypanosoma cruzi Leishmania major
Media Clip 25.4 Euglenids Human disease Sleeping sickness Chagas disease Leishmaniasis
Life12e.com/mc25.4 Insect vector Tsetse fly Assassin bugs (many Sand fly
The euglenids have diverse nutri- species)
tional requirements. Many species Vaccine or effective cure None None None
are always heterotrophic. Other spe- Strategy for survival Changes surface recognition Causes changes in Reduces
cies, including species of Euglena, molecules frequently surface recognition effectiveness of
molecules on host cell macrophage hosts
are fully autotrophic in sunlight,
using chloroplasts to synthesize Site in human body Bloodstream; in final stages, Enters cells, especially Enters cells, primarily
attacks nerve tissue muscle cells macrophages
organic compounds through pho-
Approximate number of 7,000 11,000 63,000
tosynthesis. When kept in the
deaths per year
dark, these euglenids lose their

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 KEY CONCEPT 25.2 Major Lineages of Eukaryotes Diversified in the Precambrian

Amoeba proteus these amoebas produce casings by gluing sand grains together
(Figure 25.16). Other testate amoebas have shells secreted by
the organism itself.

PLASMODIAL SLIME MOLDS If the nucleus of an amoeba be-

gan rapid mitotic division, accompanied by a tremendous increase
in cytoplasm and organelles but no cytokinesis, the resulting or-

© Wim van Egmond/Science Source

ganism would resemble the multinucleate mass of a plasmodial
slime mold. During its vegetative (feeding, nonreproductive) stage,
a plasmodial slime mold is a wall-less mass of cytoplasm with
numerous diploid nuclei. This mass streams very slowly over its
substrate in a remarkable network of strands called a plasmodium
(Figure 25.17A). The plasmodium is an example of a coenocyte:
Pseudopods many nuclei enclosed in a single cell membrane. The outer cy-
toplasm of the plasmodium (closest to the environment) is nor-
mally less fluid than the interior cytoplasm and thus provides
some structural rigidity.
Plasmodial slime molds provide a dramatic example of move-
ment by cytoplasmic streaming. The outer cytoplasm of the plas-
© Wim van Egmond/Science Source

modium becomes more fluid in places, and cytoplasm rushes

into those areas, stretching the plasmodium. This streaming
reverses its direction every few minutes as cytoplasm rushes
into a new area and drains away from a previous one, moving
the plasmodium over its substrate. Sometimes an entire wave
of plasmodium moves across a surface, leaving strands behind.
Microfilaments and a contractile protein called myxomyosin in-
120 µm teract to produce the streaming movement. As it moves, the plas-
Figure 25.15 An Amoeba in Motion The flowing pseudopods modium engulfs food particles by endocytosis—predominantly
of this “chaos amoeba” (a lobosean) are constantly changing shape bacteria, yeasts, spores of fungi, and other small organisms as
as the amoeba moves and feeds. well as decaying animal and plant remains.
Media Clip 25.5 Amoeboid Movement A plasmodial slime mold can grow almost indefinitely in its
Life12e.com/mc25.5 plasmodial stage as long as the food supply is adequate and other
conditions, such as moisture and pH, are favorable. If conditions
Nebela collaris
become unfavorable, however, one of two things can happen. First,
the plasmodium can form an irregular mass of hardened cell-like
components. This resting structure rapidly becomes a plasmo-
dium again when favorable conditions are restored.
Shell (test) Alternatively, the plasmodium can transform itself into spore-
made of bearing fruiting structures (Figure 25.17B). These stalked or
sand grains
branched structures rise from heaped masses of plasmodium.
They derive their rigidity from walls that form and thicken be-
Cell
tween their nuclei. The diploid nuclei of the plasmodium divide
membrane by meiosis as the fruiting structure develops. One or more knobs,
of amoeba called sporangia, develop on the end of the stalk. Within a sporan-
gium, haploid nuclei become surrounded by walls to form spores.
© Wim van Egmond/Science Source

Eventually, as the fruiting structure dries, it sheds its spores.

The spores germinate into wall-less, haploid cells called
swarm cells, which can either divide mitotically to produce
Pseudopods more haploid swarm cells or function as gametes. Swarm cells
of amoeba can live as separate individual cells that move by means of fla-
gella or pseudopods, or they can become walled and resistant
resting cysts when conditions are unfavorable; when conditions
18 µm improve again, the cysts release swarm cells. Two swarm cells
Figure 25.16 Life in a Glass House This testate amoeba has can also fuse to form a diploid zygote, which divides by mitosis
built a lightbulb-shaped shell, or test, by gluing sand grains together. (but without a wall forming between the nuclei) and thus forms
Its pseudopods extend through the single aperture in the test. a new coenocytic plasmodium.

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

(A) A plasmodium (B) Fruiting structures of a

When conditions become unfavorable, the
plasmodial slime mold
cellular slime molds form fruiting structures,
as do their plasmodial counterparts. The in-
dividual myxamoebas aggregate into a mass
called a slug or pseudoplasmodium. Unlike

© Matt Meadows/Science Source

the true plasmodium of the plasmodial slime
molds, this structure is not simply a giant sheet
of cytoplasm with many nuclei; the individual
myxamoebas within the slug retain their cell
membranes and therefore their identity.

David McIntyre
A slug may migrate over a substrate for sev-
30 mm eral hours before becoming motionless and
reorganizing to construct a delicate, stalked
1.5 mm
fruiting structure. Cells at the top of the fruit-
Figure 25.17 A Plasmodial Slime Mold (A) The plasmodial form of the slime mold ing structure develop into thick-walled spores,
Hemitrichia serpula covers rocks, decaying logs, and other objects as it engulfs bacteria and which are eventually released. Later, under fa-
other food items; it is also responsible for the organism’s common name of “pretzel mold.”
vorable conditions, the spores germinate, re-
(B) Fruiting structures of Leocarpus fragilis.
leasing myxamoebas.
Media Clip 25.6 Plasmodial Slime Mold Growth The cycle from myxamoebas through slug
Life12e.com/mc25.6
and spores to new myxamoebas is asexual. Cel-
lular slime molds also have a sexual cycle, in
CELLULAR SLIME MOLDS Whereas the plasmodium is the which two myxamoebas fuse. The product of this fusion develops
basic vegetative unit of the plasmodial slime molds, a single amoe- into a spherical structure that ultimately germinates, releasing
boid cell is the vegetative unit of the cellular slime molds (Figure new haploid myxamoebas.
25.18). Cells called myxamoebas, which have single haploid nuclei,
KEY CONCEPT
swarm together as they engulf bacteria and other food particles
by endocytosis and reproduce by mitosis and fission. This simple 25.2 Recap and Assess
life cycle stage, consisting of swarms of independent, isolated cells, The major lineages of eukaryotes began to diverge about 1.5 bil-
can persist indefinitely as long as food and moisture are available. lion years ago. Major groups of eukaryotes are highly diverse in
their habitat, nutrition, locomotion, and body form. Many
protists are photosynthetic autotrophs, but heterotrophic
Dictyostelium discoideum lineages have evolved repeatedly. Although most protists
The sporangium of the mature are unicellular, multicellularity has arisen independently
fruiting structure will release spores.
many times.
1. For each pair of groups below, describe how you could
Fruiting structure recognize members of the two groups and differentiate
(various stages) them from one another. Then describe features that
the two groups in each pair share.
a. Foraminiferans and radiolarians
b. Ciliates and dinoflagellates
c. Diatoms and brown algae
d. Plasmodial slime molds and cellular slime molds
2. The fossil record of eukaryotes from the Precambrian
is poor compared with that from the Cambrian and
later geological periods, even though eukaryotes
Courtesy of R. Blanton and M. Grimson

were diversifying for the last billion years of the

Precambrian. Can you think of some reasons why
the eukaryotic fossil record became more extensive
in the Cambrian?
3. Give examples of alveolates, stramenopiles, and
excavates that are important for medical or
Slug culinary reasons.
4. The red tides produced by some dinoflagellates are
0.25 mm not produced by other alveolates. But similar red tides
are produced by some diatoms, which are distantly
Figure 25.18 A Cellular Slime Mold This composite micrograph shows the related to dinoflagellates. Thus the production of red
life cycle of a Dictyostelium slime mold. tides is convergent in dinoflagellates and diatoms.
Media Clip 25.7 Cellular Slime Mold Aggregation Describe another example from the text of convergent
evolution in protists but outside alveolates.
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 KEY CONCEPT 25.3 Protists Reproduce Sexually and Asexually

The ancient origins of the major eukaryotic lineages and the adapta- reproduced groups of nearly identical organisms are known as
tion of these lineages to a wide variety of lifestyles and environments clonal lineages.
resulted in enormous protist diversity. It is not surprising, then, that Sexual reproduction among the protists takes various forms.
reproductive modes among protists are also highly diverse. In some protists, as in animals, the gametes are the only haploid
cells. In others, the zygote is the only diploid cell. In still others,
both diploid and haploid cells undergo mitosis, giving rise to al-
KEY CONCEPT Protists Reproduce Sexually ternating multicellular diploid and haploid life stages.

25.3 and Asexually Some protists reproduce without sex and

have sex without reproduction
Learning Objectives
As we noted in Key Concept 25.2, members of the genus Para-
25.3.1 Distinguish between sex (in the sense of exchange mecium are ciliates, which commonly have two types of nuclei
of genes leading to genetic recombination) and in a single cell (one macronucleus and from one to several mi-
reproduction.
cronuclei; see Figure 25.6). The micronuclei are typical eukary-
25.3.2 Predict the evolutionary consequences of otic nuclei and are essential for genetic recombination. Each
reproduction without sex for many generations.
macronucleus contains many copies of the genetic information,
packaged in units containing only a few genes each. The mac-
Although most protists engage in both asexual and sexual re- ronuclear DNA is transcribed and translated to regulate the
production, sexual reproduction has yet to be observed in some life of the cell.
groups. In some protists, as in all prokaryotes, the acts of sex and When paramecia reproduce asexually, all of the nuclei are cop-
reproduction are not directly linked. ied before the cell divides. Paramecia (and many other protists)
Several asexual reproductive processes have been observed also have an elaborate sexual behavior called conjugation, in which
among the protists: two individuals line up tightly against each other and fuse in the
• The equal splitting of one cell into two by mitosis oral groove region of the body. Nuclear material is extensively
followed by cytokinesis reorganized and exchanged over the next several hours (Figure
• The splitting of one cell into multiple (i.e., more 25.19). Each cell ends up with two haploid micronuclei, one of
than two) cells its own and one from the other cell, which fuse to form a new
diploid micronucleus. A new macronucleus develops from that
• The outgrowth of a new cell from the surface of an
micronucleus through a series of dramatic chromosomal rear-
old one (known as budding)
rangements. The exchange of nuclei is fully reciprocal: each of
• The formation of specialized cells (spores) that are capable of the two paramecia gives and receives an equal amount of DNA.
developing into new individuals (known as sporulation) The two organisms then separate and go their own ways, each
Asexual reproduction results in offspring that are genetically equipped with new combinations of alleles.
nearly identical to their parents (they differ only by new muta- Conjugation in Paramecium is a sexual process of genetic re-
tions that may arise during DNA replication). Such asexually combination, but it is not a reproductive process. Two cells begin

Micronucleus

Macronucleus

1 Two paramecia conjugate; 2 Three of the four haploid 3 The paramecia donate 4 The two micronuclei 5 The new diploid micronuclei
all but one micronucleus in micronuclei disintegrate; micronuclei to each in each cell—each divide mitotically, eventually
each cell disintegrate. The the remaining micronucleus other. The macronuclei genetically giving rise to a macronucleus
remaining micronucleus undergoes mitosis. disintegrate. different—fuse. and the appropriate number
undergoes meiosis. of micronuclei.

Figure 25.19 Conjugation in Paramecia The exchange of micro- Q: Why is conjugation considered sex without reproduction?
nuclei by two conjugating Paramecium individuals results in genetic
recombination. After conjugation, the cells separate and continue their
lives as two individuals.

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

conjugation and two cells are there at the end, so no new cells are Given the diversity of protists and of the environments in which
created. As a rule, each asexual clone of paramecia must conjugate they live, it is not surprising that they influence their environ-
periodically. Experiments have shown that if some species are not ments in numerous ways.
permitted to conjugate, the clones can live through only about 350
cell divisions before dying out. Protists Are Critical
KEY CONCEPT
Components of Many
Some protist life cycles feature
alternation of generations 25.4 Ecosystems
Alternation of generations is a type of life cycle found in many
multicellular protists, all land plants, and some fungi (see Fig- Learning Objectives
ure 26.6). A multicellular, diploid, spore-producing organism 25.4.1 Illustrate and/or describe the complex life cycle
gives rise to a multicellular, haploid, gamete-producing organ- of a parasite that involves multiple hosts.
ism. When two haploid gametes fuse, a diploid organism is pro- 25.4.2 Summarize the consequences of overproduction
duced. The haploid organism, the diploid organism, or both may of some diatoms and dinoflagellates.
also reproduce asexually. Note that alternation of generations 25.4.3 Describe some of the beneficial and harmful
is distinct from the familiar reproductive system of animals, in interactions that protists have with other species.
which the only haploid stages are unicellular gametes produced 25.4.4 Provide two examples of how protists are
by multicellular, diploid adults. important to humans.
The two alternating (spore-producing and gamete-producing)
generations differ genetically (one has diploid cells, the other hap- Some protists are food for marine animals, while others poison
loid cells), but they may or may not differ morphologically. In het- those animals. Some are packaged as nutritional supplements for
eromorphic alternation of generations, the two generations differ humans, and some are human pathogens. The remains of some
morphologically; in isomorphic alternation of generations, they do form the sands of many modern beaches, and others are a major
not. Examples of both heteromorphic and isomorphic alternation source of the oil that sometimes fouls those beaches.
of generations are found among the brown algae.
The gamete-producing generation does not produce gametes Phytoplankton are primary producers
by meiosis because the gamete-producing organism is already A single protist clade, the diatoms, performs about one-fifth of
haploid. Instead, specialized cells of the diploid spore-producing all photosynthetic carbon fixation on Earth—about the same
organism, called sporocytes, divide meiotically to produce four amount as all of Earth’s rainforests. These spectacular unicellu-
haploid spores. The spores may eventually germinate and divide lar organisms (see Figure 25.8) are the predominant component
mitotically to produce the multicellular haploid generation, which of the oceanic phytoplankton, but the phytoplankton include
then produces gametes by mitosis and cytokinesis. many other protists that also contribute heavily to global pho-
Gametes, unlike spores, can produce new organisms only by tosynthesis. Like green plants on land, these “floating photosyn-
fusing with other gametes. The fusion of two gametes produc- thesizers” are the gateway for energy from the Sun into the rest
es a diploid zygote, which then undergoes mitotic divisions to of the living world; in other words, they are primary producers.
produce a diploid organism. The diploid organism’s sporocytes These autotrophs are eaten by heterotrophs, including animals
then undergo meiosis and produce haploid spores, starting the and many other protists. Those consumers are, in turn, eaten by
cycle anew. other consumers. Most aquatic heterotrophs (with the excep-
tion of some species in the deep sea) depend on photosynthesis
KEY CONCEPT
performed by phytoplankton.
25.3 Recap and Assess
Protists reproduce both asexually and sexually, although Some microbial eukaryotes are deadly
sex occurs independently of reproduction in some species. Some microbial eukaryotes are pathogens that cause serious dis-
Some multicellular protists exhibit alternation of generations, eases in humans and other vertebrates. The best-known patho-
alternating between multicellular haploid and diploid genic protists are members of the genus Plasmodium, a highly
life stages.
specialized group of apicomplexans that spend part of their life
1. Why is conjugation between paramecia considered a
sexual process but not a reproductive process? cycle as parasites in human red blood cells, where they are the
cause of malaria. In terms of the number of people affected, ma-
2. Why do you think paramecia that are not allowed to
conjugate begin to die out after about 350 rounds of laria is one of the world’s three most serious infectious diseases:
asexual reproduction? it infects more than 215 million people, and kills nearly half a
3. Although most diploid animals have haploid stages million people, each year. On average, about two people die from
(e.g., eggs and sperm), their life cycles are not malaria every minute of every day—most of them in sub-Saharan
considered examples of alternation of generations. Africa, although malaria occurs in more than 90 countries.
Why not? Mosquitoes of the genus Anopheles transmit Plasmodium to
humans. The parasites enter the human circulatory system when

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 KEY CONCEPT 25.4 Protists Are Critical Components of Many Ecosystems

an infected female Anopheles mosquito penetrates the skin in forming zygotes. The zygotes lodge in the mosquito’s gut, divide
search of blood. The parasites find their way to cells in the liver several times, and move into its salivary glands, from which they
and the lymphatic system, change their form, multiply, and reenter can be passed on to another human host. Thus Plasmodium is an
the bloodstream, where they invade red blood cells. extracellular parasite in the mosquito vector and an intracellular
The parasites multiply inside the red blood cells, which then parasite in the human host (Figure 25.20). Such an organism—
lyse (burst), releasing new swarms of parasites. These episodes that is, a parasite that requires more than one host—is said to have
of bursting red blood cells coincide with the primary symptoms a complex life cycle.
of malaria, which include fever, shivering, vomiting, joint pains, Plasmodium has proved to be a singularly difficult pathogen to
and convulsions. If another Anopheles bites the victim, the mos- attack. The complex Plasmodium life cycle is best broken by the
quito takes in Plasmodium cells along with blood. Some of the removal of stagnant water, in which mosquitoes breed. Using in-
ingested cells develop into gametes that unite in the mosquito, secticides to reduce the Anopheles population can also be effective,

(A) (B)
1 A blood-feeding female 2 Within the mosquito, male and
mosquito ingests the female gametocytes develop
Plasmodium gametocytes. into gametes, which fuse.
Male Cysts
gamete

3 The resulting zygote

enters the mosquito’s
8 Eventually, some gut wall and forms a cyst.
merozoites develop

© London School of Hygiene/SPL/Science Source

into male and Female
female gametocytes. gamete

Mosquito’s Mosquito’s
gut wall gut wall

7b …grow and
divide, and lyse
the cells. They
can reinfect the
liver, producing
new generations.
Events in mosquito 4 The zygote gives rise 170 µm
Events in human to sporozoites that
invade the salivary gland.
Mosquito's
salivary gland
Red
blood
cell

5 The mosquito injects

7a Merozoites also sporozoites into a human’s
invade red blood blood when it feeds. The
cells,… sporozoites then take up
residence in the liver.
Human
liver cell

6 Sporozoites penetrate
liver cells and develop
into merozoites.

Figure 25.20 Life Cycle of the Malarial Parasite (A) Like many par- stomach wall of a mosquito. Invasive sporozoites will hatch from
asitic species, the apicomplexan Plasmodium falciparum has a complex the cysts and be transmitted to a human, in whom the parasite
life cycle, part of which is spent in mosquitoes of the genus Anopheles causes malaria.
and part in humans. The sexual phase (gamete fusion) of this life cycle View in Achieve
takes place in the insect, and the zygote is the only diploid stage. Animation 25.3 Life Cycle of the Malarial Parasite
(B) Encysted Plasmodium zygotes (artificially colored blue) cover the

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

but the benefits must be weighed against the ecological, economic, radiolarian can make use of the carbon compounds produced by
and health risks posed by the insecticides themselves. its photosynthetic guest, and the guest may in turn make use of
Even some of the phytoplankton that are such important metabolites made by the host or receive physical protection. In
primary producers can be deadly, as described in this chapter’s some cases, the guest is exploited for its photosynthetic products
opening story. Some diatoms and dinoflagellates reproduce in while receiving little or no benefit itself.
enormous numbers when environmental conditions are favor- Dinoflagellates are also common endosymbionts and can be
able for their growth. In the resulting red tides, the concentra- found in both animals and other protists. Most, but not all, di-
tion of dinoflagellates may reach 60 million per liter of ocean noflagellate endosymbionts are photosynthetic. Some dinoflagel-
water and produce potent nerve toxins that harm or kill many lates live endosymbiotically in the cells of corals, contributing the
vertebrates, especially fishes. products of their photosynthesis to the partnership. Their impor-
tance to the corals is demonstrated when the dinoflagellates die or
Some microbial eukaryotes are endosymbionts are expelled by the corals as a result of changing environmental
Endosymbiosis is common among the microbial eukaryotes, many conditions such as rising water temperatures or increased water
of which live within the cells of animals. Many radiolarians harbor turbidity. This phenomenon is known as coral bleaching. Unless
photosynthetic endosymbionts (see Figure 25.12A). As a result, the corals can acquire new endosymbionts, they are ultimately
these radiolarians, which are not photosynthetic themselves, ap- damaged or destroyed as a result of their reduced food supply
pear greenish or golden, depending on the type of endosymbiont (Investigating Life: Can Corals Reacquire Dinoflagellate Endo-
they contain. This arrangement is often mutually beneficial: the symbionts Lost to Bleaching?).

▶InvestigatingLIFE Can Corals Reacquire Dinoflagellate

Endosymbionts Lost to Bleaching?
Experiment 70 Experimental (exposed to strain B211)
Original Paper: C. L. Lewis and M. A. Coffroth. 2004. The 60 Control (not exposed to strain B211)
Mean number of Symbiodinium cells

acquisition of exogenous algal symbionts by an octocoral 50

per coral polyp (thousands)

after bleaching. Science 304: 1490–1492.

40 Six weeks after return to light,
Some corals lose their chief nutritional source when their both groups showed increases
30
photosynthetic endosymbionts die, often as a result of in number of symbionts present.
changing environmental conditions. This experiment by 20 DNA analysis showed that strain
Cynthia Lewis and Mary Alice Coffroth investigated the 10 B211 symbionts were present in
After 12 weeks in
ability of corals to acquire new endosymbionts after the experimental group.
dark, 0%–1% of

HYPOTHESIS▸ Bleached corals can acquire new

bleaching. the photosynthetic
3 endosymbionts

METHOD▸
photo-synthetic endosymbionts from their environment. remained.
2
1
1. Count numbers of Symbiodinium, a photosynthetic
0
dinoflagellate, living symbiotically in samples of a Pre-bleach Post-bleach Week 3 Week 6
coral (Briareum sp.). (original state)
2. Stimulate bleaching by maintaining all Briareum Pre-bleach Post-bleach
colonies in darkness for 12 weeks.
Both photos courtesy of M. A. Coffroth

3. After 12 weeks of darkness, count numbers of

and C. Lewis, University at Buffalo

Symbiodinium in the coral samples; then return all

colonies to light.
4. In some of the bleached colonies (the experimental
group), introduce Symbiodinium strain B211—
dinoflagellates that contain a unique molecular marker.
Do not expose the others (the control group) to strain

RESULTS▸ Error bars show 95% confidence limits for CONCLUSION▸ Corals can acquire new endosymbionts from
B211. Maintain both groups in the light for 6 weeks.

the means. their environment following bleaching.

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 KEY CONCEPT 25.4 Protists Are Critical Components of Many Ecosystems

We rely on the remains of ancient marine protists Other ancient marine protists have also contributed to today’s
Diatoms are lovely to look at, but their importance to us goes far world. Some foraminiferans, as we have seen, secrete shells of cal-
beyond aesthetics, and even beyond their role as primary produc- cium carbonate. After they reproduce (by mitosis and cytokinesis),
ers. Diatoms store oil as an energy reserve and to keep themselves the daughter cells abandon the parent shell and make new shells of
afloat at the correct depth in the ocean. Over millions of years, their own. The discarded shells of ancient foraminiferans make up
diatoms have died and sunk to the ocean floor, where they have extensive limestone deposits in various parts of the world, forming a
undergone chemical changes. In this way, they have become a layer hundreds to thousands of meters deep over millions of square
major source of petroleum and natural gas, two of our most im- kilometers of ocean bottom. Foraminiferan shells also make up
portant energy supplies and political concerns. much of the sand of some beaches. A single gram of such sand may
Because the silica-containing cell walls of dead diatoms resist contain as many as 50,000 foraminiferan shells and shell fragments.
decomposition, some sedimentary rocks are composed almost The shells of individual foraminiferans are easily preserved as
entirely of diatom skeletons that sank to the seafloor over time. fossils in marine sediments. Each geological period has a distinc-
Diatomaceous earth, which is obtained from such rocks, has many tive assemblage of foraminiferan species. Because the shells of
industrial uses, such as insulation, filtration, and metal polishing. foraminiferan species have distinctive shapes (see Figure 25.11)
It has also been used as an “Earth-friendly” insecticide that clogs and because they are so abundant, the remains of foraminiferans
the tracheae (breathing structures) of insects. are especially valuable in classifying and dating sedimentary rocks.
In addition, analyses of the chemical makeup of foraminiferan
shells can be used to estimate the global temperatures prevalent
at the time when the shells were formed.
KEY CONCEPT
Work with the Data 25.4 Recap and Assess
The data in the table below come from DNA analyses of
Symbiodinium strains found in the experimental and control Protists have many effects, both positive and negative, on their
colonies of corals (Briareum) before and after bleaching. environment. Some species are important primary producers,
Symbiodinium strain B211 (which was not present before many are endosymbionts, and some are pathogens. Protists are
bleaching) was introduced to the experimental colonies after among the most important producers of fossil fuels, and they
are important components of sedimentary rocks.
bleaching. Use these data to answer the questions below.
1. What is the role of female Anopheles mosquitoes in the
Symbiodinium strain present (% of colonies) transmission of malaria?
2. Explain the roles of dinoflagellates in the two very different
Non-B211 B211 Nonea Colony died
phenomena of coral bleaching and red tides.
EXPERIMENTAL COLONIES (STRAIN B211 ADDED) 3. What are two ways in which diatoms are important to
Pre-bleach 100 0 0 0 human society?
Post-bleach 58 0 42 0
Week 3 0 92 0 8 The next six chapters will explore the major evolutionary radia-
Week 6 8 58 8 25 tions of multicellular eukaryotes, along with the protist ancestors
CONTROL COLONIES (NO STRAIN B211) from which they arose. Chapters 26 and 27 will describe the origin
Pre-bleach 100 0 0 0
and diversification of plants, Chapter 28 will present the fungi,
Post-bleach 67 0 33 0
and Chapters 29–31 will provide a brief overview of the animals.
Week 3 67 0 33 0
Week 6 67 0 17 17

QUESTIONS▸
a
Colonies remained alive, but no Symbiodinium were detected.

1. Are new strains of Symbiodinium taken up only by coral

colonies that have lost all their original endosymbionts?
2. Does the acquisition of a new Symbiodinium strain
always result in survival of a recovering Briareum colony?
3. In week 3, only strain B211 was detected in the experi-
mental colonies, but in week 6, non-B211 Symbiodinium
were detected in 8% of the experimental colonies. Can
you suggest an explanation for this observation?
Go to Achieve for a companion
Data in Depth exercise.

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 CHAPTER 25 The Origin and Diversification of Eukaryotes

▶InvestigatingLIFE

QA
and
Red tides are harmful, but how are
dinoflagellates beneficial to marine ecosystems?
The opening of this chapter discussed harmful red tides, or blooms
of dinoflagellates. But not all dinoflagellate blooms produce prob-
lems for other species. Dinoflagellates are important components
of many ecosystems, as we have seen throughout this chapter. Pho-

© Per-Andre Hoffmann/Alamy Stock Photo

tosynthetic dinoflagellates also produce much of the atmospheric
oxygen that most animals need to survive.
As we saw in Investigating Life: Can Corals Reacquire Dinoflagellate
Endosymbionts Lost to Bleaching?, corals and many other species
depend on symbiotic dinoflagellates for food. In addition, as photo-
synthetic organisms, free-living planktonic dinoflagellates are among
the most important primary producers in aquatic food webs. They
are a major component of the phytoplankton and provide an impor-
tant food source for many species. Figure 25.21 Light Up the Sea Bioluminescent dinoflagellates
flash as an outrigger disturbs the ocean surface off the island of Bali.
Future directions
Some dinoflagellates produce a beautiful bioluminescence (Figure signal. However, recent research shows another function of biolu-
25.21). Unlike the bioluminescent bacteria described at the start minescence in toxic species of dinoflagellates. These toxic species
of Chapter 24, however, dinoflagellates cannot generate a steady produce a much lower intensity bioluminescence which may function
bioluminescence, but instead produce bright flashes of light when as a warning of their toxicity to potential predators. So depending
disturbed, as people who swim in the ocean at night in certain re- on the intensity and other details of the flash in different species,
gions often observe. What function do these flashes serve? Many the bioluminescence can serve either to attract secondary preda-
light-emitting dinoflagellates are preyed on by other species, such tors or to warn off primary predators. Research on communication
as small crustaceans. When the dinoflagellates produce a bright by bioluminescence continues as a rich field of investigation.
flash, it functions like a “burglar alarm,” and attracts secondary
Media Clip 25.8 Flashing Dinoflagellate
predators of the crustaceans. Experiments have shown that crusta-
Life12e.com/mc25.8
ceans reduce their feeding on dinoflagellates when they flash this

VISUAL SUMMARY 25
© Don Paulson/PureStock/AGE Fotostock

You should be able to relate each summary to the adjacent figures. If you go to this visual summary in Achieve,
you can follow links to figures, animations, activities, and simulations that will help you consolidate the material.

KEY CONCEPT

25.1 Eukaryotes Acquired Features from Both Archaea and Bacteria Go to Animation 25.1

■ The term protist does not describe a formal taxonomic ■ Mitochondria evolved by endosymbiosis with a
group. It is shorthand for “all eukaryotes that are not plants, proteobacterium.
animals, or fungi.” ■ Primary endosymbiosis of a eukaryote and a
■ Early events in the evolution of the eukaryotic cell included cyanobacterium gave rise to the first chloroplasts.
the loss of the firm cell wall and infolding of the cell Secondary endosymbiosis and tertiary endosymbiosis
membrane. Such infolding led to segregation of the genetic between chloroplast-containing eukaryotes and other
material in a membrane-enclosed nucleus. eukaryotes gave rise to the distinctive chloroplasts of
euglenids, dinoflagellates, and other groups.

25_Life12e_Ch 25.indd 19 12/3/19 4:10 PM

 VISUAL SUMMARY 25
Questions Figure 25.1 Figure 25.2
1. Are all protists small in size (microbial)? Primary endosymbiosis
Eukaryote Cyanobacterium
Cell wall
2. What is the phylogenetic evidence that
DNA
mitochondria and chloroplasts arose
through endosymbiosis?

Chloroplast

Secondary endosymbiosis
Host eukaryotic cell Chloroplast-
containing
eukaryotic cell

Flagellum
Mitochondrion
Chloroplast Nucleus

KEY CONCEPT Major Lineages of Eukaryotes

25.2 Diversified in the Precambrian Go to Activity 25.1 and Animation 25.2

■ Most eukaryotes can be placed in one of eight major ■ The amoebozoans move by means of lobe-shaped pseu-
clades that diverged about 1.5 billion years ago: alveolates, dopods and include both unicellular and multicellular
stramenopiles, rhizarians, excavates, plants, amoebozoans, members; examples are the plasmodial slime molds and
fungi, and animals. cellular slime molds.
■ Alveolates are unicellular organisms with sacs (alveoli) Questions
beneath their cell membranes. Alveolate clades include the
1. Name five of the major groups of eukaryotes in which multi-
marine dinoflagellates, the parasitic apicomplexans, and
cellularity has arisen independently.
the diverse, highly motile ciliates.
■
2. Which five of the major groups of eukaryotes contain nu-
Stramenopiles typically have two flagella of unequal length,
merous photosynthetic species?
the longer one bearing rows of tubular hairs. Among the
stramenopiles are the unicellular
diatoms; the multicellular brown
Figure 25.3
algae; and the nonphotosynthetic
Alveolates
oomycetes, many of which are
saprobic. Stramenopiles

■ Rhizarians are unicellular Rhizarians

and aquatic. They include the Excavates

cercozoans; the foraminiferans,
Plantae Chs. 26 and 27
which secrete shells of calcium
carbonate; and the radiolarians, Amoebozoans

which have thin, stiff pseudopods Fungi Ch. 28

and glassy endoskeletons.
■
Choanoflagellates
Ophisthokonts Chs.
The unicellular excavates include 29–31
Animals
parasitic as well as free-living
species, some of which have lost Precambrian Paleozoic Mesozoic Cenozoic
typical mitochondria. to 1.5 1.4 1.3 1.2 1.1 1.0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0
4.5 bya Time (bya)

25_Life12e_Ch 25.indd 20 12/3/19 4:10 PM

 VISUAL SUMMARY 25
KEY CONCEPT

25.3 Protists Reproduce Sexually and Asexually

■ Asexual reproduction gives rise to clonal lineages The alternating generations may be heteromorphic
of organisms. or isomorphic.
■ Conjugation in Paramecium is a sexual process but Questions
not a reproductive one.
1. Explain how some species can recombine their genomes
■ Alternation of generations, which involves cycles sexually, yet reproduce asexually.
between a multicellular diploid stage and a multicellular
2. Why is animal reproduction, which involves haploid gametes
haploid stage, is a feature of many multicellular protist life
produced by diploid organisms, not considered alternation
cycles (as well as those of some fungi and all land plants).
of generations?
Figure 25.19
Micronucleus

Macronucleus

KEY CONCEPT

25.4 Protists Are Critical Components of Many Ecosystems Go to Animation 25.3

■ Diatoms are responsible for about one-fifth of the photo- Questions

synthetic carbon fixation on Earth. They and other members 1. Describe two distinct problems for human populations that
of the phytoplankton are important primary producers in are caused by protists.
the marine environment. Ancient diatoms are a major source
2. What are two important benefits to humans from protists?
of today’s petroleum and natural gas deposits.
■ Some protists are pathogens of humans and other
vertebrates.
■ Figure 25.20 Male
Endosymbiotic relationships gamete
are common among microbial
protists and often benefit both Cysts
the endosymbionts and their

© London School of Hygiene/SPL/Science Source

Female
protist or animal partners. gamete

Mosquito’s
gut wall

Mosquito’s
gut wall

Events in mosquito
Events in human

Mosquito's
salivary gland 170 µm
Red
blood
cell

Human
liver cell

Go to Achieve for the eBook, LearningCurve, animations, activities,

simulations, and additional resources and assignments.

25_Life12e_Ch 25.indd 21 12/3/19 4:10 PM