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Chapter 28 Protista
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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 Movie
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Protista- single or colonies of eukaryotic cells (Ameoba, Paramecium)
Evolved from the Archae approx. 1.5 billion years ago Polyphyletic group- protists arose by way of more than one ancestral group Represents separate evolutionary lineages Plant like b/c autotrophic (produce their own food) Animal-Like b/c they are heterotrophic (feed upon other organisms) Monday you will study three phyla and animal like protists.
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Animal-Like Protists
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Cladogram of Protozoa Relationships
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Cladogram of Protozoa Relationships Fig. 8.22 8-14
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Animal-Like Protists: The Protozoa
Unicellular and Colonial Eukaryotes Study representatives of the following three Protista phyla: Phylum Sarcomastigophora Subphylum Mastigophora Subphylum Sarcodina Phylum Apicomplexa Phylum Ciliophora
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Figure 8.3
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These amazing organisms
Belong to the diverse kingdoms of mostly single-celled eukaryotes informally known as protists 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
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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
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Protist habitats are also diverse in habitat
And including freshwater and marine species Figure 28.2a–d 100 m 4 cm 500 m The freshwater ciliate Stentor, a unicellular protozoan (LM) Ceratium tripos, a unicellular marine dinoflagellate (LM) Delesseria sanguinea, a multicellular marine red alga Spirogyra, a filamentous freshwater green alga (inset LM) (a) (b) (c) (d) Reproduction and life cycles Are also highly varied among protists, with both sexual and asexual species
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A sample of protist diversity
Table 28.1
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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 Endosymbiosis in Eukaryotic Evolution
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Diversity of plastids produced by secondary endosymbiosis
Cyanobacterium Heterotrophic eukaryote Primary endosymbiosis Red algae Green algae Secondary Plastid Dinoflagellates Apicomplexans Ciliates Stramenopiles Euglenids Chlorarachniophytes Alveolates Figure 28.3
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Plasmodial slime molds
Concept 28.2: Diplomonads and parabasalids have modified mitochondria A tentative phylogeny of eukaryotes Divides eukaryotes into many clades Rhodophyta Chlorophyta Diplomonadida Euglenozoa Animalia Parabasala Plantae Cercozoa Radiolaria Fungi Alveolata Stramenopila Amoebozoa (Opisthokonta) (Viridiplantae) Fungi Plants Euglenids Ciliates Parabasalids Dinoflagellates Oomycetes Diatoms Golden algae Brown algae Radiolarians Entamoebas Metazoans Red algae Diplomonads Kinetoplastids Apicomplexans Foraminiferans Gymnamoebas Chlorophytes Chlorarachniophytes Cellular slime molds Choanoflagellates Charophyceans Plasmodial slime molds Figure 28.4 Ancestral eukaryote
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Diplomonads and parabasalids
Are adapted to anaerobic environments Lack plastids Have mitochondria that lack DNA, an electron transport chain, or citric-acid cycle enzymes Plastid-responsible for photosynthesis, storage of products like starch and for the synthesis of many classes of molecules such as fatty acids and
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(a) Giardia intestinalis, a diplomonad (colorized SEM)
Diplomonads Diplomonads Have two nuclei and multiple flagella Figure 28.5a 5 µm (a) Giardia intestinalis, a diplomonad (colorized SEM)
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(b) Trichomonas vaginalis, a parabasalid (colorized SEM)
Parabasalids Parabasalids include trichomonads Which move by means of flagella and an undulating part of the plasma membrane Flagella Undulating membrane 5 µm Figure 28.5b (b) Trichomonas vaginalis, a parabasalid (colorized SEM)
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Euglenozoa is a diverse clade that includes
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
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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 µm Crystalline rod Ring of microtubules Figure 28.6
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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
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The parasitic kinetoplastid Trypanosoma
Causes sleeping sickness in humans Figure 28.7 9 m
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Life Cycle of Trypanosoma Brucei
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Life Cycle of Trypanosoma Brucei Fig. 8.9 Other Mastigophora Zoomastigophora- Trypanosoma, Trichonympha, and Trichomonas Trichonympha- Mastigophora Symbionts- Termite gut 8-6
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Figure 8.8 (b)
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Phylum Sarcomastigophora
Chars: Flagella, pseudopodia, or both; single type of nucleus; no spores formed. Subphylum Masigophora Chars: One or more Flagella Autotrophic (cl. Phytomastigophora) Heterotrophic (cl. Zoomastigophora) or both; Reproduction usually by fission
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Subphylum Mastigophora (cl. Phytomastigophora)
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Structure of Euglena Fig. 8.7 Subphylum Mastigophora (cl. Phytomastigophora) 8-4
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Euglenids Euglenids Have one or two flagella that emerge from a pocket at one end of the cell Store the glucose polymer paramylon Figure 28.8 Long flagellum Short flagellum Nucleus Plasma membrane Paramylon granule Chloroplast Contractile vacuole 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 Eyespot: pigmented organelle that functions as a light shield, allowing light from only a certain direction to strike the light detector Pellicle: protein bands beneath the plasma membrane that provide strength and flexibility (Euglena lacks a cell wall) Euglena (LM) 5 µm Freshwater phytomastigophoran Ponds and slow moving streams
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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 Figure 28.9 Flagellum Alveoli 0.2 µm
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Dinoflagellates Dinoflagellates
Are a diverse group of aquatic photoautotrophs and heterotrophs Are abundant components of both marine and freshwater phytoplankton
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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 Figure 28.10 3 µm Flagella
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Figure 8.6 Phylum Dinozoa: Dinoflagellates
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Rapid growth of some dinoflagellates
Is responsible for causing “red tides,” which can be toxic to humans
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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
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Phylum Apicomplexa Chars: All parasites
Apical complex used for penetrating host cells Lack cilia and flagella, except in certain reproductive stages Coccidians or apicomplexans are named based upon the presence of apical complex
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Most important Coccidians are members of the class Sporozoea
Chars: intracellular parasites of animals Form spores or oocysts following sexual reproduction Complex life cycle that involve both vertebrate and invertebrate hosts Example- Plasmodium the sporozoan that causes malaria.
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Most apicomplexans have intricate life cycles
With both sexual and asexual stages that often require two or more different host species for completion An infected Anopheles mosquito bites a person, injecting Plasmodium sporozoites in its saliva. 1 The sporozoites enter the person’s 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). 2 Figure 28.11 Inside mosquito Inside human Sporozoites (n) Oocyst MEIOSIS Liver Liver cell Merozoite Red blood cells Gametocytes FERTILIZATION Gametes Zygote (2n) Key Haploid (n) Diploid (2n) cell Apex 0.5 µm An oocyst develops from the zygote in the wall of the mosquito’s gut. The oocyst releases thousands of sporozoites, which migrate to the mosquito’s salivary gland. 7 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. 3 Gametes form from gametocytes. Fertilization occurs in the mosquito’s digestive tract, and a zygote forms. The zygote is the only diploid stage in the life cycle. 6 Another Anopheles mosquito bites the infected person and picks up Plasmodium gametocytes along with blood. 5 Some merozoites form gametocytes. 4
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Life Cycle of Plasmodium
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Life Cycle of Plasmodium Fig. 8.15 8-10
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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
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Phylum Ciliophora Chars: Cilia, macronuclei, and micronuclei usually present Ciliates are the largest most complex and diverse group of the protozoans Nearly occupy all aquatic habitats Some are symbiotic Reproduction can be asexual through fission or sexual through conjugation
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Example of a Ciliophora: Paramecium
Common freshwater ciliate Observe live sample using methylcellulose solution Other Ciliophora: Colpidium, Vorticella and Stentor
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FEEDING, WASTE REMOVAL, AND WATER BALANCE
Exploring structure and function in a ciliate 50 µm 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 Food vacuoles combine with lysosomes. As the food is digested, the vacuoles follow a looping path through the cell. Thousands of cilia cover the surface of Paramecium. Micronucleus Macronucleus The micronuclei Function during conjugation, a sexual process that produces genetic variation Conjugation is separate from reproduction Which generally occurs by binary fission 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. Figure 28.12
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CONJUGATION AND REPRODUCTION
8 7 2 MICRONUCLEAR FUSION Diploid micronucleus Haploid micronucleus MEIOSIS Compatible mates Key Conjugation Reproduction Macronucleus Two cells of compatible mating strains align side by side and partially fuse. 1 Meiosis of micronuclei produces four haploid micronuclei in each cell. 2 3 Three micronuclei in each cell disintegrate. The remaining micro- nucleus in each cell divides by mitosis. The cells swap one micronucleus. 4 The original macro- nucleus disintegrates. Four micronuclei become macronuclei, while the other four remain micronuclei. 8 The cells separate. 5 Micronuclei fuse, forming a diploid micronucleus. 6 Two rounds of cytokinesis partition one macronucleus and one micronucleus into each of four daughter cells. 9 Three rounds of mitosis without cytokinesis produce eight micronuclei. 7
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Asexual Reproduction in Protozoa - ciliophora
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Binary Fission of Ciliated Stentor
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Stentor ciliophora The trumpet animalcule Stentor is one of the largest unicellular organisms. Most of the time it lives attached to a surface. With it's cell stretched Stentor feeds on bacteria and other small creatures using a crown of fused cilia (hairlike structures). But when disturbed it uses it's cilia to locomote. As an aid to reach a large size, sometimes two millimetres long, Stentor has within it's cell a string of many nucleï. The algae live in symbiosis with the Stentor, ie the algae and Stentor mutually benefit from the close association. The algae uses photosynthesis to convert Stentor's waste products to useful nutrients.
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Concept 28.5: Stramenopiles have “hairy” and smooth flagella
The clade Stramenopila Includes several groups of heterotrophs as well as certain groups of algae
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Most stramenopiles Have a “hairy” flagellum paired with a “smooth” flagellum Smooth flagellum Hairy 5 µm Figure 28.13
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germinate, growing into
The life cycle of a water mold Encysted zoospores land on a substrate and germinate, growing into a tufted body of hyphae. 1 Several days later, the hyphae begin to form sexual structures. 2 Meiosis produces eggs within oogonia (singular, oogonium). 3 On separate branches of the same or different individuals, meiosis produces several haploid sperm nuclei contained within antheridial hyphae. 4 Figure 28.14 Cyst Zoospore (2n) ASEXUAL REPRODUCTION Zoosporangium Germ tube Zygote germination FERTILIZATION SEXUAL Zygotes (oospores) MEIOSIS Oogonium Egg nucleus (n) Antheridial hypha with sperm nuclei Key Haploid (n) Diploid (2n) Each zoospor- angium produces about 30 biflagellated zoospores asexually. 9 The ends of hyphae form tubular zoosporangia. 8 The zygotes germinate and form hyphae, and the cycle is completed. 7 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. 5 A dormant period follows, during which the oogonium wall usually disintegrates. 6
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Diatoms Diatoms are unicellular algae
With a unique two-part, glass-like wall of hydrated silica Figure 28.15 3 µm
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Diatoms are a major component of phytoplankton
And are highly diverse Figure 28.16 50 µm
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Most golden algae are unicellular
But some are colonial Figure 28.17 25 µm Brown algae, or phaeophytes Are the largest and most complex algae Are all multicellular, and most are marine The cells of golden algae Are typically biflagellated, with both flagella attached near one end of the cell
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Brown algae Include many of the species commonly called seaweeds Seaweeds Have the most complex multicellular anatomy of all algae Figure 28.18 Blade Stipe Holdfast
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Kelps, or giant seaweeds
Live in deep parts of the ocean Figure 28.19
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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
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Foraminiferans (Forams)
Foraminiferans, or forams Are named for their porous, generally multichambered shells, called tests Figure 28.22 20 µm
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Pseudopodia extend through the pores in the test
Foram tests in marine sediments Form an extensive fossil record
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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
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The pseudopodia of radiolarians, known as axopodia
Radiate from the central body Figure 28.23 200 µm Axopodia
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Concept 28.7: Amoebozoans have lobe-shaped pseudopodia Amoebozoans
Are amoeba that have lobe-shaped, rather than threadlike, pseudopodia Include gymnamoebas, entamoebas, and slime molds
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Gymnamoebas Gymnamoebas
Are common unicellular amoebozoans in soil as well as freshwater and marine environments
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Most gymnamoebas are heterotrophic
And actively seek and consume bacteria and other protists Figure 28.24 Pseudopodia 40 µm
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Entamoebas Entamoebas Entamoeba histolytica
Are parasites of vertebrates and some invertebrates Entamoeba histolytica Causes amebic dysentery in humans Chars: Pseudopodia, Flagella occasionally present (in developmental stages. The Amoebas Ameobas- common freshwater protist Lives on the bottom of ponds
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Variations in Pseudopodia
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Variations in Pseudopodia Fig. 8.10 8-7
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Subphylum Sarcodina: Superclass Rhizopoda, Class Lobosea
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Subphylum Sarcodina: Superclass Rhizopoda, Class Lobosea Fig. 8.11b 8-8
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Other Sarcodina-“Not naked” sarcodines
Arcella, Difflugia, and Actinospaerium and marine radiolarians and foraminifera form test. Test can be formed from sand grains, calcium carbonate and silica
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Freshwater Amoeba (Difflugia Oblongata)
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Freshwater Amoeba (Difflugia Oblongata) Fig. 8.12 8-9
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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
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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
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Most chlorophytes Other 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
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Chlorophytes include Unicellular, colonial, and multicellular forms
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). (a) Caulerpa, an inter- tidal chlorophyte. The branched fila- ments lack cross-walls and thus are multi- nucleate. In effect, the thallus is one huge “supercell.” (b) 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. (c) 20 µm 50 µm Figure 28.30a–c Phytomstigophoran- Volvox
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Volvox, A Colonial Flagellate
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Volvox, A Colonial Flagellate Fig. 8.8 8-5
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Most chlorophytes have complex life cycles
With both sexual and asexual reproductive stages 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. 7 In Chlamydomonas, mature cells are haploid and contain a single cup-shaped chloroplast (see TEM at left). 1 In response to a shortage of nutrients, drying of the pond, or some other stress, cells develop into gametes. 2 Gametes of opposite mating types (designated + and –) pair off and cling together. Fusion of the gametes (syngamy) forms a diploid zygote. 3 Figure 28.31 Flagella Cell wall Nucleus Regions of single chloroplast Zoospores ASEXUAL REPRODUCTION Mature cell (n) SYNGAMY SEXUAL Zygote (2n) MEIOSIS 1 µm Key Haploid (n) Diploid (2n) + The zygote secretes a durable coat that protects the cell against harsh conditions. 4 When a mature cell repro- duces asexually, it resorbs its flagella and then undergoes two rounds of mitosis, forming four cells (more in some species). 6 After a dormant period, meiosis produces four haploid individuals (two of each mating type) that emerge from the coat and develop into mature cells. 5
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