CHAPTER

Diversification of Eukaryotes
27

Learning Objectives: Students should be able to...

27.1 Summarize the medical and ecological importance of protists.
27.1.1 Explain the impact of protists on human health.
27.1.2 Describe the ecological importance of protists.
27.2 Evaluate evolutionary relationships among protists, and explain how this group is
paraphyletic.
27.2.1 Explain the role of microscopy in the study of protist structure.
27.2.2 Evaluate phylogenies to identify relationships among Eukarya lineages.
27.2.3 Explain how new taxa are discovered.
27.3 Describe the life history innovations that evolved in protists.
27.3.1 Explain how endosymbiosis lead to the origin of mitochondria and chloroplasts.
27.3.2 Explain how the plasma membrane may have played a key role in the origin of
nuclei.
27.3.3 Describe the ways by which protists obtain nutrients.
27.3.4 Explain how protists move.
27.3.5 Explain the role of “alternation of generations” in the life cycle of protists.
27.4 Describe the key features of the major lineages of eukaryotes.
27.4.1 Describe the characteristics of the following taxa: Amoebozoa, Excavata, Plantae,
Rhizaria, Alveolata, Stramenopila.

Lecture Outline
 Protists are eukaryotes.
 Fundamental features of eukaryotes:
 The nuclear envelope is a synapomorphy that defines the domain Eukarya.
 Most eukaryotic cells are larger than prokaryotes, have many organelles, and have an
extensive system of structural proteins called the cytoskeleton.
 Multicellularity has evolved multiple times in eukaryotes.
 Eukaryotes reproduce either asexually via mitosis or sexually via meiosis.
 Protists are all the eukaryotes that are not fungi, green plants, or animals.
 Protists are a collection of lineages which surrounded by water most of the time.
(Figs. 27.1, 27.2)
I. Why Do Biologists Study Protists?
A. Impacts on human health and welfare
1. The Irish potato famine in 1845 was caused by a protist, Phytophthora infestans.
a. One million people (out of a population of 9 million) died, and millions of others
emigrated to other countries.
2. Malaria
a. Malaria is the world’s worst chronic health problem, affecting 200 million people
worldwide each year and killing 600,000 people each year.
b. It is caused by five species of the protist genus Plasmodium.
c. Plasmodium cells enter the bloodstream via a mosquito bite, infects liver cells and
(later) red blood cells, and destroy the cells. (Fig. 27.3)
d. Plasmodium is probably the best studied of all the protists, but we still do not have
effective measures to control it.
(1) Plasmodium has evolved resistance to many drugs.
(2) Efforts to develop a vaccine have not been successful.
(3) Mosquitoes have continually evolved resistance to insecticides.
e. Many other protists also cause human disease. (Table 27.1)
3. Harmful algal blooms
a. Algal blooms occur when a unicellular aquatic species experiences rapid population
growth.
b. Many harmful algal blooms are caused by photosynthetic protists called dinoflagel-
lates, which contain toxins. (Fig. 27.4)
c. High levels of toxins build up in shellfish that filter the protists out of the water, and
people who eat the shellfish can then get sick or die.
B. Ecological importance of protists
1. Protists are extremely abundant in their habitats. For example, there are more than 500
flagellated protists per milliliter of pond water and 60 thousand dinoflagellates per
milliliter of seawater.
2. Protists play a key role in aquatic food chains as primary producers.
a. Diatoms and other protists that drift in the ocean are called plankton.
3. Could protists help limit global climate change?
a. Phytoplankton in the ocean takes up large quantities of carbon dioxide and often
carry it to the bottom of the ocean when they die, all as part of the global carbon
cycle. (Fig. 27.5)
b. Fertilizing the ocean with iron speeds up the carbon cycle and might increase the
amount of carbon taken into the marine carbon sink.
c. Once at the bottom of the ocean, they may enter one of two carbon sinks:
(1) Sedimentary rocks—mostly made of calcium carbonate
(2) Petroleum—oil formation tends to begin with accumulation of dead bacteria,
archaea, and protists
II. How Do Biologists Study Protists?
A. Microscopy: studying cell structure
1. Detailed studies of cell structure revealed that many protists have a characteristic over-
all form, organelles with distinctive features, or both. (BioSkills 9)
 a. For example, special flagella with tiny hairlike projections, known as stra-
menopiles, have been identified on the protist that caused the potato famine and on
some types of brown algae. (Fig. 27.6)
2. Seven major groups of eukaryotes have been identified in this way, based on diagnos-
tic morphological features (synapomorphies) that either protect or support the cell or
aid in reproduction or feeding. (Table 27.2)
III. What Themes Occur in the Diversification of Protists?
A. What morphological innovations evolved in protists?
1. All eukaryotes alive today have (1) either mitochondria or mitochondrial genes,
(2) a nucleus and endomembrane system, and (3) a cytoskeleton.
2. Early eukaryotes were probably single-celled organisms with mitochondria, a nucleus
and endomembrane system, and a cytoskeleton, but no cell wall.
a. Students should be able to map the origin of the nuclear envelope and the
eukaryotic flagellum in Figure 27.6.
3. Endosymbiosis and the origin of the mitochondrion
a. The endosymbiosis theory (Lynn Margulis, 1970s) proposes that mitochondria
evolved from an aerobic bacterium that was engulfed by an anaerobic eukaryotic
cell. (Fig. 27.8)
(1) This became a mutually beneficial symbiosis: The host supplied the bacterium
with protection and carbon compounds, and the bacterium produced much more
ATP than the host could produce on its own.
b. Many lines of evidence now support this theory.
(1) Mitochondria are similar in size to -proteobacteria.
(2) They divide independently of the host cell, and they divide by fission, as
bacteria do.
(3) They have their own ribosomes and synthesize their own proteins, and the
ribosomes are similar in many ways to bacterial ribosomes.
(4) They have double membranes, as would be expected if they were engulfed by
another cell.
(5) They have their own chromosomes, which are circular and similar to bacterial
chromosomes.
c. The most conclusive evidence is that mitochondrial genes are very closely related to
the genes from -proteobacteria
4. Endosymbiosis and the origin of the chloroplasts
a. All photosynthetic protists have chloroplasts.
b. An extension of the endosymbiotic theory contends that the eukaryotic chloroplast
originated when a protist engulfed a cyanobacterium.
c. Many lines of evidence support this:
(1) Chloroplasts have the same list of bacteria-like characteristics presented earlier
for mitochondria.
(2) They contain circular DNA containing genes extremely similar to genes found
in various species of cyanobacteria.
(3) The photosynthetic organelle of glaucophyte algae have an outer layer
containing the same peptidoglycan found in cyanobacteria.
 (4) There are many examples of endosymbiotic cyanobacteria living inside cells of
protists or animals today.
d. To explain the fact that many protists have chloroplasts containing multiple layers
of membrane, researchers proposed the process of secondary endosymbiosis—an
organism engulfs a photosynthetic eukaryotic cell and retains the chloroplast.
(Fig. 27.9)
e. Figure 27.10 shows where primary and secondary endosymbiosis occurred in the
phylogenetic tree of eukaryotes.
f. Students should be able to assess and explain why the primary and secondary
endosymbiosis events introduced in this chapter represent the most massive
lateral gene transfers in the history of life, in terms of number of genes moved at
once. (Chapter 20, lateral gene transfer)
5. The nuclear envelope
a. The nuclear envelope likely arose as infoldings of the plasma membrane that also
gave rise to the endoplasmic reticulum. (Fig. 27.11)
b. Evidence: Infoldings of the plasma membrane occur in some extant bacteria, and
the nuclear envelope and ER of today’s eukaryotes are continuous.
c. Students should be able to explain why the above observations support the
infolding hypothesis.
d. The nuclear envelope allowed for the separation of transcription and translation.
e. This enabled alternative splicing and gave early eukaryotes more ways to control
gene expression.
6. Structures for support and protection
a. The diversification of protists has been associated with the evolution of innovative
structures for support and protection:
(1) Glasslike shells in a box-and-lid arrangement (diatoms) (Fig. 27.12a)
(2) A cell wall made of cellulose plates (dinoflagellates) (Fig. 27.12b)
(3) Chambers of calcium carbonate (foraminiferans) (Fig. 27.12c)
(4) Coverings of tiny pebbles (other foraminiferans and some amoebae)
(5) An internal support rod (parabasalids)
(6) A collection of protein strips under the plasma membrane (euglenids)
(Fig. 27.12d)
(7) Saclike structures (alveolates)
7. Multicellularity
a. Multicellular organisms contain more than one cell and have cells that are special-
ized for different functions and express different genes.
b. The vast majority of multicellular species are eukaryotes.
c. Multicellularity evolved independently in several eukaryotic lineages: green plants,
fungi, animals, brown algae, slime molds, and red algae.
B. How do protists obtain food?
1. Protists have three ways of feeding: ingesting packets of food, absorption of molecules

directly from the environment, and photosynthesis.

 a. Some species of protists use multiple feeding methods, and all three methods may
occur within a single lineage. Example: alveolates
2. Ingestive feeding
a. The large cell size of protists enabled some of them to develop a unique feeding
strategy: ingesting other organisms (ingestive feeding).
b. Ingestion can occur by eating live or dead organisms or scavenging loose bits of
organic debris.
c. Protists that feed by phagocytosis typically lack a cell wall and can move their
plasma membranes around their prey using pseudopodia. (Fig. 27.13a)
d. Some ingestive feeders sit and wait for prey to come by or have cilia that move the
environment by them. (Fig. 27.13b)
3. Absorptive feeding
a. Some absorptive-feeding protists are decomposers and feed on detritus (dead
organic matter).
b. Others live inside other organisms.
(1) Absorptive feeders that damage their hosts are called parasites.
4. Photosynthesis
a. Organisms that produce their own food through photosynthesis are called
autotrophs.
b. Organisms that ingest or absorb food are called heterotrophs.
C. How do protists move?
1. Amoeboid motion is a sliding movement produced by extensions of the cell, called
pseudopodia, into which the cytoplasm streams. (Fig. 27.14)
2. Other protists move using cilia or flagella.
a. Flagella and cilia are identical in structure; each is composed of nine sets of doublet
microtubules and two central, single microtubules.
b. Flagella are long and usually single or paired (Fig. 27.15a). Cilia are short and
numerous. (Fig. 27.15b)
3. Even closely related protists can use very different forms of locomotion.
D. How do protists reproduce?
1. Sexual reproduction evolved in protists and is one of the most significant evolutionary
innovations of the eukaryotes.
a. Asexual reproduction occurs via mitosis and cell division (protists) or fission
(bacteria, archaea). Offspring are genetically identical to the parents.
b. Sexual reproduction occurs via meiosis and fusion of gametes. Offspring are
genetically different from the parents.
2. Sexual versus asexual reproduction
a. Genetically variable offspring have a better chance of surviving in a changing
environment.
b. Pathogens and parasites are highly variable aspects of the environment because they
evolve quickly. Thus, sexual reproduction may primarily be an adaptation to fight
disease and parasites.
3. Students should be able to evaluate and explain whether it is consistent with the
observation that many protists undergo meiosis only when food is scarce or popula-
tion density is high.
IV. Key Lineages of Protists (Table 27.3)
 There are seven major lineages in the Eukarya (six are described in this section).
 Each of the seven has at least one distinctive morphological characteristic but has also di-
versified greatly in other ways.
 There have been multiple independent origins of parasitism, photosynthesis, and
multicellularity.
A. Amoebozoa
1. Amoebozoa lack cell walls, feed by engulfing, and move by amoeboid movement with
large, lobelike pseudopodia.
a. Students should be able to mark the origin of this lineage’s amoeboid form in
Figure 27.7.
2. They include lobose amoebae, cellular slime molds, and plasmodial slime molds.
3. Plasmodial slime molds (Table 27.3)
a. Traits: All form a large, multinucleated “supercell,” feed on decaying vegetation,
move by cell crawling, and use sexual reproduction when food is scarce.
b. Human and ecological impacts: They are important decomposers in forests.
c. Students should be able to mark the origin of the supercell and the origin of this
lineage’s amoeboid form in Figure 27.7.
B. Excavata
1. Excavata cells have a feeding groove on one side and have lost their mitochondria.
a. Students should be able to mark the loss of mitochondria in Figure 27.7.
2. They include parabasilids, diplomonads, and euglenids.
3. Parabasalids, Diplomonadids, and Euglenids
a. Traits: Parabasalids have distinctive rods of cross-linked microtubules. Diplomona-
dids possess two nuclei, each with four flagella. Euglenoids have a system of
interlocking protein structure to support the cell and some possess light sensitive
eyespot. Nearly all members of Excavata reproduce only asexually.
b. Human and ecological impacts: The parabasalid Trichomonas and the diplomoinid,
Giardia intestinalis are widespread human pathogens. Euglenids are important
components of freshwater food chains.
C. Plantae
1. Plantae have chloroplasts and perform photosynthesis.
2. The term Plantae is increasingly used to include glaucophyte algae (“blue-green
algae”), red algae, green algae, and land plants.
3. Students should be able to mark the origin of the plant chloroplast in Figure 27.7.
4. Red algae
a. Traits: They are primarily marine; their cell walls have cellulose; and they are pho-
tosynthetic, many with red pigments that allow them to live in deeper water. Most
are multicellular. They are the only type of algae that lack flagella. Alternation of
generations is common.
b. Human and ecological impacts: They stabilize coral reefs. The nori seaweed
(Porphyra) used to wrap sushi is a red alga.
D. Rhizaria
1. Rhizaria are single-celled amoebae that lack cell walls; move by amoeboid motion;
and produce long, slender pseudopodia.
a. Students should be able to mark the origin of this lineage’s amoeboid form in
Figure 27.7. Students should also be able to explain whether amoeboid form
evolved just once or independently in separate protist lineages.
2. They include foraminifera, actinopods, chlorarachniophytes, and many others.
3. Foraminiferans (Table 27.3)
a. Traits: All are marine, have multiple nuclei per cell, produce shells stiffened by cal-
cium carbonate, feed by engulfing, and have no locomotion; they just float in water.
They use both asexual and sexual reproduction.
b. Human and ecological impacts: They are abundant in marine sediments, where they
form deposits that become chalk, limestone, and marble.
E. Alveolata
1. Alveolates are unicellular and have small sacs called alveoli just under the cell
membrane.
2. They include ciliates, dinoflagellates, apicomplexans, and others.
3. Students should be able to mark the origin of alveoli in Figure 27.7.
4. Ciliates, Dinoflagellates, and Apicomplexans (Fig. 27.18)
a. Traits: Ciliate cells have two nuclei: macronucleus and micronucleus. Dinoflagel-
late species each has a distinct shape some maintained by cellulose plates. Some
species are bioluminescent. Apicomplexans lack flagella or cilia.
b. Human and ecological impacts: Ciliates are abundant marine plankton and live in
the guts of grazing animals, helping them digest cellulose. Photosynthetic dinoflag-
ellates are important primary producers in marine environments. Two important
apicomplexans are Plasmodium (malaria) and Toxoplasma.
F. Stramenopila (Heterokonta)
1. Stramenopila have a unique “hairy” flagellum at some point in their lifecycle.
2. They include water molds, diatoms, and brown algae.
3. Students should be able to mark the origin of “hairy” flagellum in Figure 27.7.
4. Water molds, Diatoms, and Brown Algae
a. Traits: Some water molds are unicellular; others are multicellular with long, branch-
ing filaments called hyphae. Most diatoms are unicellular, often in a box-and-lid
arrangement. Brown algae are multicellular with bodies consisting of leaflike
blades, a stipe, and a holdfast. Most species of stramenopiles exhibit diploid
dominant lifecycles.
b. Human and ecological impacts: Water molds are important decomposers in aquatic
ecosystems. Diatoms are considered the most primary producer in marine and fresh-
water systems. Brown algae form kelp forests in cold marine habitats.
 Chapter Vocabulary
To emphasize the functional meanings of these terms, the list is organized by topic rather than by first mention in
the chapter. It includes terms that may have been introduced in earlier chapters but are important to the current
chapter as well. It also includes terms other than those highlighted in bold type in the chapter text.

absorptive feeding engulfing Opisthokonta

alternation of gen- euglenid parabasalid
erations Eukarya paraphyletic
Alveolata eukaryote parasite
amoeba (plural: Excavata petroleum (oil)
amoebae) fertilization picoplankton
amoeboid motion fission plankton
Amoebozoa flagellum Plantae
Apicomplexa flagellum (plural: flagella) plasmodial slime mold
asexual reproduc- food chain Plasmodium
tion Foraminifera primary endosymbiosis
Bikonta gamete primary producer
bioluminescence gametophyte protist
blade glaucophyte algae pseudopodia
brown algae global carbon cycle red algae
cellular slime mold global warming Rhizaria
Ciliata green algae secondary endosymbiosis
cilium (plural: harmful algal bloom sedimentary rocks
cilia) holdfast sessile
conjugation hypha (plural: hyphae) sexual reproduction
cytoskeleton ingestive feeding shell
decomposer life cycle spore
detritus light microscopy sporophyte
diatom macronucleus stipe
diatomaceous earth malaria Stramenopila
Dinoflagellata meiosis symbiosis
dinoflagellate micronucleus synapomorphy
diplomonad mitochondria transmission electron mi-
direct sequencing mitosis croscopy
endomembrane system multicellularity Unikonta
endosymbiosis theory nuclear envelope water mold