Life: the Science of Biology

Lecture Notebook to accompany

Sinauer Associates, Inc. MacMillan

Copyright © 2014 Sinauer Associates, Inc. Cover photograph © Alex Mustard/naturepl.com. This document may not be modified or distributed (either electronically or on paper) without the permission of the publisher, with the following exception: Individual users may enter their own notes into this document and may print it for their own personal use. The Origin and Diversification of 0027 Eukaryotes

27.1 A Hypothetical Sequence for the Evolution of the Eukaryotic Cell (Page 550) 1 The protective Cell wall cell wall was lost. DNA

2 Infolding of the plasma membrane added surface area without increasing the cell’s volume.

3 Cytoskeleton (microfilament and microtubules) formed.

4 Internal membranes studded with ribosomes formed.

5 As regions of the infolded plasma membrane enclosed the cell’s DNA, a precursor of a nucleus formed. 6 Microtubules from the cytoskeleton formed the eukaryotic flagellum, 7 Early digestive vacuoles enabling propulsion. evolved into lysosomes using enzymes from the early endoplasmic reticulum.

8 Mitochondria formed through endosymbiosis with a proteobacterium.

9 Endosymbiosis with cyanobacteria led to the development of chloroplasts.

Flagellum

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(A) Primary endosymbiosis Eukaryote Cyanobacterium

Cyanobacterium outer membrane

Peptidoglycan

Cyanobacterium inner membrane

Host cell nucleus Chloroplast

Peptidoglycan has been lost except in glaucophytes.

Chloroplast- (B) Secondary endosymbiosis containing eukaryotic cell

Host eukaryotic cell

Host membrane (from endocytosis) encloses the engulfed cell.

A trace of the engulfed cell’s nucleus is retained in some groups.

The engulfed cell’s plasma membrane (white) has been lost in euglenids and dinoflagellates.

27.2 Endosymbiotic Events in the Evolution of Chloroplasts (Page 552)

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.02 05-24-12

Alveolates

Stramenopiles

Rhizaria

Excavates

Plantae Chs. 28 and 29

Amoebozoans

Fungi Ch. 30

Ophisthokonts Choanoflagellates Chs. 31–33 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 Billions of years ago 27.3 Precambrian Divergence of Major Eukaryote Groups (Page 553)

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.03 05-24-12

Dinoflagellates Alveolates

Apicomplexans

Ciliates

Stramenopiles

Rhizaria

In-Text Art (Page 553)

Alveolates

Brown algae Stramenopiles

Diatoms

Oomycetes

Rhizaria

Alveolates

Stramenopiles

Cercozoans Rhizaria

Foraminiferans

Radiolarians

Excavates Diplomonads

Parabasalids

Heteroloboseans

Euglenids

Kinetoplastids

Amoebozoans Loboseans

Plasmodial slime molds

Peridinium Cellularsp. slime molds

Equatorial LIFE The Science of Biology 10E Sadava groove Sinauer Associates Morales Studio Figure 27.Cladograms 05-24-12

Longitudinal groove

27.4 A Dinoflagellate (Page 554)

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.03 05-24-12 Chapter 27 | The Origin and Diversification of Eukaryotes 6

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

Cilia 10 µm Bands of cilia 7 µm Oral groove Rows of fused cilia 25 µm 27.5 Diversity among the Ciliates (Page 554)

POL Hillis Sinauer Associates Figure 20.05 Date 07-30-10

The macronucleus controls the cell’s activities. Micronuclei function in genetic recombination.

Contractile vacuole

Alveoli

Cilia Digestive vacuole Oral groove

Anal pore Trichocyst Fibrils Pellicle Alveolus Cilium

27.6 Anatomy of Paramecium (Page 555)

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.06 05-24-12 Chapter 27 | The Origin and Diversification of Eukaryotes 7 INVESTIGATINGLIFE 27.7 The Role of Vacuoles in Ciliate Digestion

HYPOTHESIS The digestive vacuoles of Paramecium produce an acidic environment that allows the organism to digest food particles. Method 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 and sequence of color (i.e., acid level) changes.

Results

1 A digestive vacuole forms around yeast cells.

2 The change in color shows that the interior Stained vacuole has become yeast cells acidic.

Oral groove

3 As products of digestion move into the cytosol, the pH increases in the vacuole (the dye becomes red again). 4 Red-stained (basic) waste material is expelled.

CONCLUSION Some ciliates acidify digestive vacuoles to assist in the breakdown of food. Go to BioPortal for discussion and relevant links for all INVESTIGATINGLIFE figures.

(Page 555)

LIFE The Science of Biology 10e Sinauer Associates Figure 2707 Date 05-21-12

Apicomplexans

Ciliates

Stramenopiles

Rhizaria

Chapter 27 | The Origin and Diversification of Eukaryotes 8

Alveolates

Brown algae Stramenopiles

Diatoms

Oomycetes

Rhizaria

In-Text Art (Page 555)

Alveolates

Stramenopiles

Cercozoans Rhizaria

Foraminiferans

Radiolarians

Excavates Diplomonads

Parabasalids

Heteroloboseans

Euglenids

Kinetoplastids

Amoebozoans Loboseans

Plasmodial slime molds

Cellular slime molds

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.Cladograms 05-24-12

Diatoms display ...or bilateral 25 µm either radial (circular) (left-right) symmetry... symmetry.

27.8 Diatom Diversity (Page 556)

(A) Himanthalia elongata (B) Postelsia palmiformis

27.9 Brown Algae (Page 556) Holdfasts

LIFE 10E Sinauer Associates SaprolegniaFigure 27.09 sp.

27.10 An Oomycete (Page 557) 3 mm

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.10 Date 04-20-09 Dinoflagellates Alveolates

Apicomplexans

Ciliates

Stramenopiles

Rhizaria

Alveolates

Brown algae Stramenopiles

Diatoms

Oomycetes

Rhizaria

Chapter 27 | The Origin and Diversification of Eukaryotes 10

Alveolates

Stramenopiles

Cercozoans Rhizaria

Foraminiferans

Radiolarians In-Text Art (Page 557)

Excavates Diplomonads

Parabasalids

Heteroloboseans

Euglenids

Kinetoplastids

Amoebozoans Loboseans

Plasmodial slime molds

Cellular slime molds

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.Cladograms 05-24-12 1 mm 27.11 Building Blocks of Limestone (Page 557)

(A) (B)

Astrolithium sp. Hexacontium sp. 250 µm 50 µm LIFE The Science of Biology 10E Sadava 27.12 Radiolarians Exhibit Distinctive Pseudopods and Radial Symmetry (Page 557) Sinauer Associates Morales Studio Figure 27.11 Date 04-20-09 © 2014 Sinauer Associates, Inc.

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.11 Date 04-20-09 Dinoflagellates Alveolates

Apicomplexans

Ciliates

Stramenopiles

Rhizaria

Alveolates

Brown algae Stramenopiles

Diatoms

Oomycetes

Rhizaria

Alveolates

Stramenopiles

Cercozoans Rhizaria

Foraminiferans

Radiolarians

Chapter 27 | The Origin and Diversification of Eukaryotes 11

Excavates Diplomonads

Parabasalids

Heteroloboseans

Euglenids

Kinetoplastids In-Text Art (Page 558)

Amoebozoans Loboseans

Plasmodial slime molds

Cellular slime molds

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales(A) Giardia Studio sp. Figure 27.Cladograms 05-24-12

2.5 µm

(B) Trichomonas vaginalis

2.5 µm

27.13 Some Excavate Groups Lack Mitochondria (Page 558)

LIFE The Science of Biology 9E Sadava Sinauer Associates Morales Studio Figure 27.24 Date 04-20-09 Chapter 27 | The Origin and Diversification of Eukaryotes 12

Photosynthetic chloroplasts are prominent features in a typical Euglena cell.

Flagella Nucleus

Pigment shield

Contractile Stored polysaccharides Photoreceptor vacuole from photosynthesis 27.14 A Photosynthetic Euglenid (Page 559)

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.14 Date 05-24-12

TablE27.1 Three Pathogenic Trypanosomes Trypanosoma brucei Trypanosoma cruzi Leishmania major Human disease Sleeping sickness Chagas disease Leishmaniasis Insect vector Tsetse fly Assassin bugs (many species) Sand fly Vaccine or effective cure None None None Strategy for survival Changes surface recognition Causes changes in surface recognition Reduces effectiveness of macrophage molecules frequently molecules on host cell hosts Site in human body Bloodstream; in final stages, Enters cells, especially muscle cells Enters cells, primarily macrophages attacks nerve tissue Approximate number of 50,000 45,000 60,000 deaths per year

(Page 559)

Apicomplexans

Ciliates

Stramenopiles

Rhizaria

Alveolates

Brown algae Stramenopiles

Diatoms

Oomycetes

Rhizaria

Alveolates

Stramenopiles

Cercozoans Rhizaria

Foraminiferans

Radiolarians

Excavates Diplomonads

Parabasalids

Heteroloboseans

Euglenids

Kinetoplastids Chapter 27 | The Origin and Diversification of Eukaryotes 13

Amoebozoans Loboseans

Plasmodial slime molds

Cellular slime molds

In-Text Art (Page 559)

Chaos carolinensis LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.Cladograms 05-24-12

Pseudopods

Nebela collaris

120 µm

27.15 An Amoeba in Motion (Page 559) Shell (test) made of sand grains

Plasma membrane of amoeba

Pseudopods of amoeba

18 µm 27.16 Life in a Glass House (Page 560)

LIFE The Science of Biology 9E Sadava Sinauer Associates Morales Studio Figure 2707 Date 05-12-09 Chapter 27 | The Origin and Diversification of Eukaryotes 14

(A)

30 mm (B)

27.17 A Plasmodial Slime Mold 1.5 mm (Page 560)

POL Hillis Sinauer Associates Figure 20.16 Date 07-30-10

The sporangium of the mature fruiting structure Dictyostelium discoideum will release spores.

Fruiting structure (various stages)

Slug

0.25 mm 27.18 A Cellular Slime Mold (Page 561)

LIFE The Science of Biology 9E Sadava Sinauer Associates Morales Studio Figure 27.29 Date 04-20-09 Chapter 27 | The Origin and Diversification of Eukaryotes 15

Macronucleus

Micronucleus

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.

27.19 Conjugation in Paramecia (Page 562)

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.19 Date 05-24.12

(A) START

8 Eventually, some 1 A blood-feeding female merozoites develop mosquito ingests the into male and Plasmodium gametocytes. 7 Merozoites also female gametocytes. invade red blood cells, grow and Male divide, and lyse the gamete cells. They can 2 Within the mosquito, reinfect the liver, male and female producing new gametocytes develop generations. into gametes, which fuse.

Red blood cell Female gamete 3 The resulting zygote enters the mosquito’s gut wall and forms a cyst.

Events in human Events in mosquito

Mosquito's gut wall

4 The zygote gives rise 6 Sporozoites penetrate to sporozoites that liver cells and develop invade the salivary gland. into merozoites. (B) Human liver cell Mosquito's salivary gland

5 The mosquito injects sporozoites into a human’s blood when it feeds. Cysts 27.20 Life Cycle of the Malarial Parasite (Page 564)

Mosquito's gut wall

170 µm

LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 27.20 05-24-12

HYPOTHESIS Bleached corals can acquire new photosynthetic endosymbionts from their environment. Method 1. Count numbers of Symbiodinium, a photosynthetic dinoflagellate, living symbiotically in samples of a coral (Briareum sp.). 2. Stimulate bleaching by maintaining all Briareum colonies in darkness for 12 weeks. 3. After 12 weeks of darkness, count numbers of 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 B211. Maintain both groups in the light for 6 weeks.

Results 70 Experimental (exposed to strain B211) 60 Control (not exposed to strain B211) 50

40 Six weeks after return to light, both groups showed 30 increases in number of 20 symbionts present. DNA analysis showed that strain 10 After 12 weeks in B211 symbionts were present dark, 0–1% of the in the experimental group. photosynthetic endosymbionts 3 remained. per coral polyp (thousands) 2 Mean number of Symbiodinium cells 1 0 Pre-bleach Post-bleach Week 3 Week 6 (original state)

Pre-bleach Post-bleach

CONCLUSION Corals can acquire new endosymbionts from their environment following bleaching. Go to BioPortal for discussion and relevant links for all INVESTIGATINGLIFE figures.

(Page 565) Life10e27.21 5/21/12