28 - Protists

Chapter 28
Protists
PowerPoint Lectures for Biology, Seventh Edition

Neil Campbell and Jane Reece
Lectures by Chris Romero

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Overview: A World in a Drop of Water Even a low-power microscope

Can reveal an astonishing menagerie of organisms in a drop of pond water
Figure 28.1
50 m
These amazing organisms

Belong to the diverse kingdoms of mostly single-celled eukaryotes informally known as protists
Advances in eukaryotic systematics

Have caused the classification of protists to change significantly
Concept 28.1: Protists are an extremely diverse assortment of eukaryotes Protists are more diverse than all other eukaryotes
And are no longer classified in a single kingdom
Most protists are unicellular

And some are colonial or multicellular
Protists, the most nutritionally diverse of all eukaryotes, include

Photoautotrophs, which contain chloroplasts Heterotrophs, which absorb organic molecules or ingest larger food particles
Mixotrophs, which combine photosynthesis and heterotrophic nutrition
Protist habitats are also diverse in habitat

And including freshwater and marine species
(a) The freshwater ciliate Stentor, a unicellular protozoan (LM) 100 m 100 m
4 cm
(b) Ceratium tripos, a unicellular marine dinoflagellate (LM)
(c) Delesseria sanguinea, a multicellular marine red alga 500 m
Figure 28.2ad
(d) Spirogyra, a filamentous freshwater green alga (inset LM)
Reproduction and life cycles

Are also highly varied among protists, with both sexual and asexual species
A sample of protist diversity
Table 28.1
Endosymbiosis in Eukaryotic Evolution There is now considerable evidence

That much of protist diversity has its origins in endosymbiosis
The plastid-bearing lineage of protists

Evolved into red algae and green algae
On several occasions during eukaryotic evolution

Red algae and green algae underwent secondary endosymbiosis, in which they themselves were ingested
Diversity of plastids produced by secondary endosymbiosis

Plastid Dinoflagellates Alveolates Apicomplexans
Secondary endosymbiosis
Cyanobacterium
Red algae
Ciliates
Primary endosymbiosis
Stramenopiles
Heterotrophic eukaryote
Plastid
Euglenids
Secondary endosymbiosis
Green algae
Figure 28.3
Chlorarachniophytes
Figure 28.4
Diplomonads Parabasalids Parabasala Kinetoplastids Euglenozoa Euglenids Dinoflagellates Apicomplexans Ciliates Oomycetes Diatoms Golden algae Brown algae Chlorarachniophytes Cercozoa Foraminiferans Radiolarians Radiolaria Gymnamoebas Entamoebas Plasmodial slime molds Cellular slime molds Fungi Choanoflagellates Metazoans Red algae Chlorophytes Charophyceans Plants Amoebozoa Ancestral eukaryote Alveolata Diplomonadida Stramenopila Fungi (Opisthokonta) Animalia Rhodophyta Chlorophyta Plantae (Viridiplantae)
A tentative phylogeny of eukaryotes

Divides eukaryotes into many clades
Concept 28.2: Diplomonads and parabasalids have modified mitochondria
Diplomonads and parabasalids

Are adapted to anaerobic environments
Lack plastids
Have mitochondria that lack DNA, an electron transport chain, or citric-acid cycle enzymes
Diplomonads Diplomonads
Have two nuclei and multiple flagella
Figure 28.5a
(a) Giardia intestinalis, a diplomonad (colorized SEM)
5 m
Parabasalids Parabasalids include trichomonads

Which move by means of flagella and an undulating part of the plasma membrane
Flagella
Undulating membrane 5 m Figure 28.5b (b) Trichomonas vaginalis, a parabasalid (colorized SEM)
Concept 28.3: Euglenozoans have flagella with a unique internal structure Euglenozoa is a diverse clade that includes
Predatory heterotrophs, photosynthetic autotrophs, and pathogenic parasites
The main feature that distinguishes protists in this clade

Is the presence of a spiral or crystalline rod of unknown function inside their flagella
Flagella 0.2 m
Crystalline rod
Figure 28.6
Ring of microtubules
Kinetoplastids Kinetoplastids
Have a single, large mitochondrion that contains an organized mass of DNA called a kinetoplast Include free-living consumers of bacteria in freshwater, marine, and moist terrestrial ecosystems
The parasitic kinetoplastid Trypanosoma

Causes sleeping sickness in humans
Figure 28.7
9 m
Euglenids Euglenids
Have one or two flagella that emerge from a pocket at one end of the cell Store the glucose polymer paramylon
Long flagellum Eyespot: pigmented organelle that functions as a light shield, allowing light from only a certain direction to strike the light detector
Short flagellum Euglena (LM) Nucleus 5 m Plasma membrane Pellicle: protein bands beneath the plasma membrane that provide strength and flexibility (Euglena lacks a cell wall)
Light detector: swelling near the base of the long flagellum; detects light that is not blocked by the eyespot; as a result, Euglena moves toward light of appropriate intensity, an important adaptation that enhances photosynthesis Contractile vacuole
Chloroplast Paramylon granule
Figure 28.8
Concept 28.4: Alveolates have sacs beneath the plasma membrane Members of the clade Alveolata
Have membrane-bounded sacs (alveoli) just under the plasma membrane
0.2 m
Flagellum
Alveoli
Figure 28.9
Dinoflagellates Dinoflagellates
Are a diverse group of aquatic photoautotrophs and heterotrophs Are abundant components of both marine and freshwater phytoplankton
Each has a characteristic shape

That in many species is reinforced by internal plates of cellulose
Two flagella
Make them spin as they move through the water
Flagella
Figure 28.10
Rapid growth of some dinoflagellates

Is responsible for causing red tides, which can be toxic to humans
Apicomplexans Apicomplexans
Are parasites of animals and some cause serious human diseases Are so named because one end, the apex, contains a complex of organelles specialized for penetrating host cells and tissues Have a nonphotosynthetic plastid, the apicoplast
Most apicomplexans have intricate life cycles

With both sexual and asexual stages that often require two or more different host species for completion
1 An infected Anopheles mosquito bites a person, injecting Plasmodium sporozoites in its saliva. 2 The sporozoites enter the persons liver cells. After several days, the sporozoites undergo multiple divisions and become merozoites, which use their apical complex to penetrate red blood cells (see TEM below). Inside mosquito Sporozoites (n) 7 An oocyst develops from the zygote in the wall of the mosquitos gut. The oocyst releases thousands of sporozoites, which migrate to the mosquitos salivary gland. Inside human Merozoite
Liver
Liver cell
Oocyst MEIOSIS Zygote (2n) Merozoite (n) Red blood cells Red blood cell
Apex 0.5 m
FERTILIZATION Gametes Key Haploid (n) Diploid (2n) 6 Gametes form from gametocytes. Fertilization occurs in the mosquitos digestive tract, and a zygote forms. The zygote is the only diploid stage in the life cycle. 5 Another Anopheles mosquito bites the infected person and picks up Plasmodium gametocytes along with blood. Gametocytes (n)
3 The merozoites divide asexually inside the red blood cells. At intervals of 48 or 72 hours (depending on the species), large numbers of merozoites break out of the blood cells, causing periodic chills and fever. Some of the merozoites infect new red blood cells.
4 Some merozoites form gametocytes.
Figure 28.11
Ciliates Ciliates, a large varied group of protists

Are named for their use of cilia to move and feed Have large macronuclei and small micronuclei
The micronuclei
Function during conjugation, a sexual process that produces genetic variation
Conjugation is separate from reproduction

Which generally occurs by binary fission
Exploring structure and function in a ciliate

FEEDING, WASTE REMOVAL, AND WATER BALANCE
Paramecium, like other freshwater protists, constantly takes in water by osmosis from the hypotonic environment. Bladderlike contractile vacuoles accumulate excess water from radial canals and periodically expel it through the plasma membrane. Contractile Vacuole Paramecium feeds mainly on bacteria. Rows of cilia along a funnel-shaped oral groove move food into the cell mouth, where the food is engulfed into food vacuoles by phagocytosis. Oral groove Cell mouth
50 m
Thousands of cilia cover the surface of Paramecium. Micronucleus Macronucleus
Food vacuoles combine with lysosomes. As the food is digested, the vacuoles follow a looping path through the cell.
Figure 28.12
The undigested contents of food vacuoles are released when the vacuoles fuse with a specialized region of the plasma membrane that functions as an anal pore.
CONJUGATION AND REPRODUCTION

1 Two cells of compatible mating strains align side by side and partially fuse. 2 Meiosis of micronuclei produces four haploid micronuclei in each cell. 3 Three micronuclei in each cell disintegrate. The remaining micronucleus in each cell divides by mitosis.
MEIOSIS
4 The cells swap one micronucleus.
Macronucleus
Compatible mates
Diploid micronucleus Diploid micronucleus
Haploid micronucleus
MICRONUCLEAR FUSION
The cells separate.
9 Two rounds of cytokinesis partition one macronucleus and one micronucleus into each of four daughter cells.
8 The original macronucleus disintegrates. Four micronuclei become macronuclei, while the other four remain micronuclei.
7 Three rounds of mitosis without cytokinesis produce eight micronuclei.
6 Micronuclei fuse, forming a diploid micronucleus.
Key Conjugation Reproduction
Concept 28.5: Stramenopiles have hairy and smooth flagella The clade Stramenopila
Includes several groups of heterotrophs as well as certain groups of algae
Most stramenopiles
Have a hairy flagellum paired with a smooth flagellum
Hairy flagellum Smooth flagellum
Figure 28.13
5 m
Oomycetes (Water Molds and Their Relatives) Oomycetes

Include water molds, white rusts, and downy mildews Were once considered fungi based on morphological studies
Most oomycetes
Are decomposers or parasites
Have filaments (hyphae) that facilitate nutrient uptake
The life cycle of a water mold

1 Encysted zoospores land on a substrate and germinate, growing into a tufted body of hyphae. 2 Several days later, the hyphae begin to form sexual structures. Germ tube Cyst 9 Each zoosporangium produces about 30 biflagellated zoospores asexually. 8 The ends of hyphae form tubular zoosporangia. MEIOSIS ASEXUAL REPRODUCTION Zoospore (2n) Oogonium Egg nucleus (n) Antheridial hypha with sperm nuclei (n) 3 Meiosis produces eggs within oogonia (singular, oogonium). 4 On separate branches of the same or different individuals, meiosis produces several haploid sperm nuclei contained within antheridial hyphae.
Zoosporangium (2n) Key

Haploid (n) Diploid (2n)
SEXUAL FERTILIZATION Zygote REPRODUCTION germination Zygotes (oospores) (2n)
7 The zygotes germinate and form hyphae, and the cycle is completed.
Figure 28.14
6 A dormant period follows, during which the oogonium wall usually disintegrates.
5 Antheridial hyphae grow like hooks around the oogonium and deposit their nuclei through fertilization tubes that lead to the eggs. Following fertilization, the zygotes (oospores) may develop resistant walls but are also protected within the wall of the oogonium.
The ecological impact of oomycetes can be significant

Phytophthora infestans causes late blight of potatoes
Diatoms Diatoms are unicellular algae

With a unique two-part, glass-like wall of hydrated silica
Figure 28.15
Diatoms are a major component of phytoplankton

And are highly diverse
Figure 28.16
50 m
Accumulations of fossilized diatom walls

Compose much of the sediments known as diatomaceous earth
Golden Algae Golden algae, or chrysophytes

Are named for their color, which results from their yellow and brown carotenoids
The cells of golden algae

Are typically biflagellated, with both flagella attached near one end of the cell
Most golden algae are unicellular

But some are colonial
25 m
Figure 28.17
Brown Algae Brown algae, or phaeophytes

Are the largest and most complex algae
Are all multicellular, and most are marine
Brown algae
Include many of the species commonly called seaweeds
Seaweeds
Have the most complex multicellular anatomy of all algae
Blade
Stipe
Figure 28.18
Holdfast
Kelps, or giant seaweeds

Live in deep parts of the ocean
Figure 28.19
Human Uses of Seaweeds Many seaweeds

Are important commodities for humans
Are harvested for food

(a) The seaweed is grown on nets in shallow coastal waters.
(b) A worker spreads the harvested seaweed on bamboo screens to dry.
Figure 28.20ac
(c) Paper-thin, glossy sheets of nori make a mineral-rich wrap for rice, seafood, and vegetables in sushi.
Alternation of Generations A variety of life cycles

Have evolved among the multicellular algae
The most complex life cycles include an alternation of generations

The alternation of multicellular haploid and diploid forms
The life cycle of the brown alga Laminaria

1 The sporophytes of this seaweed are usually found in water just below the line of the lowest tides, attached to rocks by branching holdfasts. Sporangia 2 In early spring, at the end of the main growing season, cells on the surface of the blade develop into sporangia. 3 Sporangia produce zoospores by meiosis. Sporophyte (2n) 7 The zygotes grow into new sporophytes, starting life attached to the remains of the female gametophyte.
MEIOSIS Zoospores
Developing sporophyte Zygote (2n)
Female Gametophytes (n)
Egg Mature female gametophyte (n) Key FERTILIZATION
Male
Sperm 6 Sperm fertilize the eggs.
Figure 28.21
4 The zoospores are all structurally alike, but about half of them develop into male gametophytes and half into female gametophytes. The gametophytes look nothing like the sporophytes, being short, branched filaments that grow on the surface of subtidal rocks. 5 Male gametophytes release sperm, and female gametophytes produce eggs, which remain attached to the female gametophyte. Eggs secrete a chemical signal that attracts sperm of the same species, thereby increasing the probability of fertilization in the ocean.
Concept 28.6: Cercozoans and radiolarians have threadlike pseudopodia A newly recognized clade, Cercozoa
Contains a diversity of species that are among the organisms referred to as amoebas
Amoebas were formerly defined as protists

That move and feed by means of pseudopodia
Cercozoans are distinguished from most other amoebas

By their threadlike pseudopodia
Foraminiferans (Forams) Foraminiferans, or forams

Are named for their porous, generally multichambered shells, called tests
20 m
Figure 28.22
Pseudopodia extend through the pores in the test Foram tests in marine sediments
Form an extensive fossil record
Radiolarians Radiolarians are marine protists

Whose tests are fused into one delicate piece, which is generally made of silica That phagocytose microorganisms with their pseudopodia
The pseudopodia of radiolarians, known as axopodia

Radiate from the central body
Axopodia Figure 28.23

200 m
Concept 28.7: Amoebozoans have lobeshaped pseudopodia Amoebozoans

Are amoeba that have lobe-shaped, rather than threadlike, pseudopodia Include gymnamoebas, entamoebas, and slime molds
Gymnamoebas Gymnamoebas
Are common unicellular amoebozoans in soil as well as freshwater and marine environments
Most gymnamoebas are heterotrophic

And actively seek and consume bacteria and other protists
Pseudopodia 40 m
Figure 28.24
Entamoebas Entamoebas
Are parasites of vertebrates and some invertebrates
Entamoeba histolytica
Causes amebic dysentery in humans
Slime Molds Slime molds, or mycetozoans

Were once thought to be fungi
Molecular systematics
Places slime molds in the clade Amoebozoa
Plasmodial Slime Molds Many species of plasmodial slime molds

Are brightly pigmented, usually yellow or orange
4 cm
Figure 28.25
At one point in the life cycle

They form a mass called a plasmodium
1 The feeding stage is a multinucleate plasmodium that lives on organic refuse. 2 The plasmodium takes a weblike form. 3 The plasmodium erects stalked fruiting bodies (sporangia) when conditions become harsh.
Feeding plasmodium Zygote (2n) SYNGAMY
Mature plasmodium (preparing to fruit) Young sporangium 1 mm
Amoeboid cells (n) Spores (n)
Mature sporangium Key
Flagellated cells (n) 7 The cells unite in pairs (flagellated with flagellated and amoeboid with amoeboid), forming diploid zygotes.
Germinating spore
MEIOSIS
Stalk
Figure 28.26
6 These cells are either amoeboid or flagellated; the two forms readily convert from one to the other.
5 The resistant spores disperse through the air to new locations and germinate, becoming active haploid cells when conditions are favorable.
Within the bulbous tips of the sporangia, meiosis produces haploid spores.
The plasmodium
Is undivided by membranes and contains many diploid nuclei Extends pseudopodia through decomposing material, engulfing food by phagocytosis
Cellular Slime Molds Cellular slime molds form multicellular aggregates

In which the cells remain separated by their membranes
The life cycle of Dictyostelium, a cellular slime mold

1 In the feeding 9 In a favorable environment, amoebas stage of the life emerge from the spore cycle, solitary haploid coats and begin feeding. amoebas engulf bacteria.
2 During sexual reproduction, two haploid amoebas fuse and form a zygote. 8 Spores are released. 7 Other cells crawl up the stalk and develop into spores.
600 m
SYNGAMY
Emerging Spores amoeba (n)

Solitary amoebas (feeding stage) ASEXUAL REPRODUCTION Aggregated amoebas Migrating aggregate
SEXUAL REPRODUCTION MEIOSIS Amoebas
Zygote (2n)
3 The zygote becomes a giant cell (not shown) by consuming haploid amoebas. After developing a resistant wall, the giant cell undergoes meiosis followed by several mitotic divisions. 4 The resistant wall ruptures, releasing new haploid amoebas.
Fruiting bodies
6 The aggregate migrates for a while and then stops. Some of the cells dry up after forming a stalk that supports an asexual fruiting body.
5 When food is depleted, hundreds of amoebas congregate in response to a chemical attractant and form a sluglike aggregate (photo below left). Aggregate formation is the beginning of asexual reproduction. Key
200 m
Figure 28.27
Dictyostelium discoideum
Has become an experimental model for studying the evolution of multicellularity
Concept 28.8: Red algae and green algae are the closest relatives of land plants Over a billion years ago, a heterotrophic protist acquired a cyanobacterial endosymbiont
And the photosynthetic descendants of this ancient protist evolved into red algae and green algae
Red Algae Red algae are reddish in color

Due to an accessory pigment call phycoerythrin, which masks the green of chlorophyll
Red algae
Are usually multicellular; the largest are seaweeds Are the most abundant large algae in coastal waters of the tropics
(b) Dulse (Palmaria palmata). This edible species has a leafy form. (c) A coralline alga. The cell walls of coralline algae are hardened by calcium carbonate. Some coralline algae are members of the biological communities around coral reefs.
Figure 28.28ac
(a) Bonnemaisonia hamifera. This red alga has a filamentous form.
Green Algae Green algae

Are named for their grass-green chloroplasts
Are divided into two main groups: chlorophytes and charophyceans

Are closely related to land plants
Most chlorophytes
Live in fresh water, although many are marine
Other chlorophytes
Live in damp soil, as symbionts in lichens, or in snow
Figure 28.29
Chlorophytes include
Unicellular, colonial, and multicellular forms
20 m 50 m (a) Volvox, a colonial freshwater chlorophyte. The colony is a hollow ball whose wall is composed of hundreds or thousands of biflagellated cells (see inset LM) embedded in a gelatinous matrix. The cells are usually connected by strands of cytoplasm; if isolated, these cells cannot reproduce. The large colonies seen here will eventually release the small daughter colonies within them (LM).
(b) Caulerpa, an intertidal chlorophyte. The branched filaments lack cross-walls and thus are multinucleate. In effect, the thallus is one huge supercell.
Figure 28.30ac
(c) Ulva, or sea lettuce. This edible seaweed has a multicellular thallus differentiated into leaflike blades and a rootlike holdfast that anchors the alga against turbulent waves and tides.
Most chlorophytes have complex life cycles

With both sexual and asexual reproductive stages
7 These daughter cells develop flagella and cell walls and then emerge as swimming zoospores from the wall of the parent cell that had enclosed them. The zoospores grow into mature haploid cells, completing the asexual life cycle. 1 In Chlamydomonas, mature cells are haploid and contain a single cup-shaped chloroplast (see TEM at left). 2 In response to a shortage of nutrients, drying of the pond, or some other stress, cells develop into gametes. Flagella Cell wall
1 m
Nucleus
+ +
Zoospores ASEXUAL REPRODUCTION Mature cell (n) SEXUAL REPRODUCTION
3 Gametes of opposite mating types (designated + and ) pair off and cling together. Fusion of the gametes (syngamy) forms a diploid zygote.
SYNGAMY
Regions of single chloroplast
Zygote (2n)
+
Key Haploid (n) Diploid (2n)
MEIOSIS 4 The zygote secretes a durable coat that protects the cell against harsh conditions.
6 When a mature cell reproduces asexually, it resorbs its flagella and then undergoes two rounds of mitosis, forming four cells (more in some species).
Figure 28.31
5 After a dormant period, meiosis produces four haploid individuals (two of each mating type) that emerge from the coat and develop into mature cells.

28 - Protists

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Chapter 28

PowerPoint Lectures for Biology, Seventh Edition

Lectures by Chris Romero

Overview: A World in a Drop of Water Even a low-power microscope

These amazing organisms

Advances in eukaryotic systematics

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Most protists are unicellular

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Protists, the most nutritionally diverse of all eukaryotes, include

Mixotrophs, which combine photosynthesis and heterotrophic nutrition

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Protist habitats are also diverse in habitat

(b) Ceratium tripos, a unicellular marine dinoflagellate (LM)

(c) Delesseria sanguinea, a multicellular marine red alga 500 m

(d) Spirogyra, a filamentous freshwater green alga (inset LM)

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Reproduction and life cycles

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

A sample of protist diversity

Endosymbiosis in Eukaryotic Evolution There is now considerable evidence

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The plastid-bearing lineage of protists

On several occasions during eukaryotic evolution

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Diversity of plastids produced by secondary endosymbiosis

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

A tentative phylogeny of eukaryotes

Concept 28.2: Diplomonads and parabasalids have modified mitochondria

Diplomonads and parabasalids

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

(a) Giardia intestinalis, a diplomonad (colorized SEM)

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Parabasalids Parabasalids include trichomonads

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The main feature that distinguishes protists in this clade

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The parasitic kinetoplastid Trypanosoma

Chloroplast Paramylon granule

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Each has a characteristic shape

Rapid growth of some dinoflagellates

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Most apicomplexans have intricate life cycles

4 Some merozoites form gametocytes.

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Ciliates Ciliates, a large varied group of protists

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Conjugation is separate from reproduction

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Exploring structure and function in a ciliate

Thousands of cilia cover the surface of Paramecium. Micronucleus Macronucleus

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

CONJUGATION AND REPRODUCTION

Diploid micronucleus Diploid micronucleus

The cells separate.

7 Three rounds of mitosis without cytokinesis produce eight micronuclei.

6 Micronuclei fuse, forming a diploid micronucleus.

Key Conjugation Reproduction

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings