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Extended Lecture Outline |
Chapter 30: The Evolution Of Eucaryotes |
A. PROBLEMS OF EUCARYOTIC TAXONOMY
30.1 The classification of many eucaryotes is problematic.
a. Kingdom Protista has historically included various hard-to-classify organisms, mostly microorganisms.
b. Many investigators are currently using modern molecular methods to sort out the phylogeny and taxonomy of these diverse, problematic, and often poorly known organisms.
30.2 Basal eucaryotes include three general nutritional types of organisms.
a. Protozoans can be defined as unicellular, non-photosynthetic organisms that feed like animals by ingesting bits of food (Figure 30.2).
1. Ciliates are single cells that propel themselves with fine cilia.
2. Flagellates move by means of long flagella, often only two but sometimes many.
3. Amoebas are often quite shapeless and move by extending rounded pseudopods, which they also use to engulf food.
b. Some molds are closely related to the golden-brown and yellow-green algae, and are classified among the protists.
1. Molds are osmiotrophs, generally saprobes or saprophytes, which decompose the detritus of an ecosystem and absorb nutrients from the water and soil; or they are parasites, which absorb nutrients from the cells of other organisms.
2. Saprobes produce their own extracellular digestive enzymes and pick up the released monomers, or they rely on the digestive enzymes produced by other organisms around them.
3. This general category also includes slime molds which are often brightly colored creatures that creep over wood and other decaying vegetation.
c. Most basal eucaryotes are relatively simple, undifferentiated, photosynthetic organisms known as algae (singular, alga; Figure 30.1).
1. Many common algae are grass-green organisms related to land plants and are placed in the plant kingdom for that reason.
2. Other algae are red, golden-brown, yellow-green, brown, and even colorless.
30.3 Several shared features provide guidelines to the phylogeny of eucaryotes.
a. A series of features, summarized in Table 30.1 point to the existence of three major clades among eucaryotes, what are called the green, brown, and red clades. The red clade (true Fungi) will be discussed in Chapter 31.
b. In spite of their basic similarity, the chloroplasts of various algae differ in significant ways and are important shared derived features.
1. The green-clade algae have chloroplasts with single thylakoids (Figure 30.4).
2. In the chloroplasts of these organisms, the thylakoids then occur in bundles of three (Figure 30.4).
3. The phototrophs are also remarkable in having their chloroplasts in the lumen of the rough endoplasmic reticulum, instead of the cytosol.
4. Algal chloroplasts produce food reserves that are often stored in the pyrenoids which are visible in Figure 30.4).
c. Although eucaryotic flagella and cilia all have the same basic 9 + 2 structure, some differences in their surface structures correlate with other phylogenetic indicators.
1. The two main types of flagellawhiplash and tinselare shown in Figure 30.5.
2. The green-clade algae generally have a pair of whiplash flagella, and some have more than one pair.
3. The organisms typically have one whiplash flagellum (often pointed backward) and one tinsel flagellum (often pointed forward).
4. In some groups, only the distinctive tinsel flagellum remains.
d. Metabolic pathways are generally universal, because once an adequate basic process has evolved there is no selective pressure to change it.
1. Lysine is an exception to this "rule."
2. Bacteria synthesize lysine through a metabolic pathway that includes diaminopimelic acid (DAP), which is a common component of their cell walls (Figure 30.6).
3. Most eucaryotes that can make their own lysine also use the DAP pathway, but a few groups use a different pathway that includes a-aminoadipic acid (AAA) (Figure 30.6).
B. A SURVEY OF THE BASAL EUCARYOTES
30.4 The chromists take many forms.
a. Some of the simplest chromists are single cells with golden, brownish or yellow-green chloroplasts and a pair of flagella for movement, and they are classified as golden-brown algae (phylum Chrysophyta) and yellow-green algae (phylum Xanthophyta) (Figure 30.7).
1. Single cells with no flagella are coccoid, and similar cells that stay cemented together in packets embedded in mucilage are called tetrasporine, even though the cells are rarely in groups of four and are not spores of any kind.
2. Many algae are filaments, often highly branched filaments, although this form is not common among chromists (Figure 30.8).
3. Filamentous algae may also have a siphonaceous form, which are long tubes not divided into cellular segments, so several nuclei reside in a single cytoplasm (a coenocytic condition).
4. Chromists also include rhizopodial algae or amoebas, so called because of their branching pseudopods (Figure 30.9).
5. Chrysophytes and xanthophytes are common members of both freshwater and marine plankton.
b. Chromist algae called diatoms (phylum Bacillariophyta) have magnificently sculptured shells built of hydrated silica (silicon dioxide, the chief component of glass) in an organic matrix (Figure 30.10).
1. Figure 30.11 outlines the cell division cycle of a diatom.
2. Diatoms lack flagella or cilia, but remain motile because of slits (called raphes) in each half-shell which are associated with nearby bundles of contractile fibrils.
3. The fibrils move along the raphe until they adhere to some object, then they contract and if the object is large enough the diatom can move along it by means of these fibrils.
c. Among the most interesting chromists are golden-brown to greenish-brown organisms called dinoflagellates (phylum Pyrophyta; Figure 30.12).
1. Most dinoflagellates are unicellular and distinctively shaped, covered with a cellulose armor divided into plates, and split by two grooves that contain flagella.
2. Some dinoflagellates use the energy of ATP to "light up" by phosphorescence in the ocean at night.
3. Other dinoflagellates are the cause of "red tide" which ultimately poisons fish and other vertebrates who feed on molluscs that have fed on the dinoflagellates.
d. The oomycetes (phylum Oomyceta) are simple fungi that include water molds and downy mildews (Figure 30.13).
1. Oomycetes have flagellated reproductive cells, cellulose cell walls, and some other features showing their relationship to the algae.
2. Oomycetes probably evolved from some very similar siphonaceous yellow-green algae.
3. Most oomycetes are obscure saprobes that grow in water or in moist soil, but downy mildews are plant pathogens on important crops.
e. The most conspicuous chromist algae are the brown algae (phylum Phaeophyta), mostly multicellular organisms that include the prominent brown seaweeds called kelps that dominate rocky coasts (Figure 30.14).
1. Brown algae show little or no differentiation of their tissues other than reproductive cells. The general term for such undifferentiated multicellular organisms is thallus (plural, thalli).
2. Their tissues are made of thin-walled, box-like cells called parenchyma.
3. Brown algae may be long and filamentous, finely branched, or flat and leaflike. Some kelps form large sheets and cylinders of parenchymal cells.
4. Brown algae and other seaweeds are important in the diets of some people who live near seashores.
30.5 Red algae have some primitive features.
a. Nearly 4,000 species of red algae constitute the phylum Rhodophyta.
1. They are mostly marine and are abundant in warm tropical coastal waters, although some species live in fresh water and soil.
2. The red algae range from single cells to lacy, interwoven filaments and broad sheets, marked by distinctive red-purple colors from their phycobilin pigments (Figure 30.15).
3. The phycobilin pigments are built into distinctive particles (phycobilisomes) covering their photosynthetic membranes (Figure 30.16).
b. Red-algae cells are surrounded by heavy walls with an inner layer of cellulose and an outer layer of pectic materials, like the pectins of fruit.
1. Coralline red algae, whose walls contain calcium deposits, are important components of coral reefs.
2. After cell division in red algae, the new wall does not close off completely but leaves a connection, like a plasmodesma in plant cells (Figure 30.17).
3. There are many similarities between the complex life cycles of many red algae and those of rust fungi (Figure 30.18).
30.6 Several types of protozoans probably evolved from algae.
a. Many of the colorless amoebas, phylum Sarcodina, undoubtedly arose from the similar rhizopodial algae of the Chromista.
1. These heterogeneous organisms, though united by their pseudopods, are probably not all closely related.
2. Amoeba proteus is a common freshwater example of a protozoan with probable algal roots (Figure 30.19).
3. Figure 30.19 shows an amoeba with formidable-looking pseudopods that resemble flagella.
4. Foraminiferans secrete calcium carbonate shells that eventually turn into limestone and chalk on the ocean floor.
5. Because the discarded shells of foraminiferans are so distinctive and well-preserved, geologists use them extensively to determine the ages of sediments.
b. The heliozoans and radiolarians produce symmetrical skeletons of silica, suggesting a relationship with the diatoms and other chromist algae that metabolize silica so well.
c. The sporozoa are all parasites with complex life cycles, a diverse group now divided into at least two phyla, Apicomplexa and Cnidosporidia.
1. The Apicomplexa include Plasmodium species, the agents of malaria in humans.
2. Plasmodium goes through several stages as it cycles from person to person through Anopheles mosquitoes (Figure 30.20).
d. Ciliates (phylum Ciliophora) are a large, diverse group of organisms that propel themselves with the coordinated movements of the cilia that cover their surfaces (Figure 30.21).
1. Connecting fibrils coordinate the whole ciliary field, so the cilia beat together and the direction of movement can be changed quickly.
2. Scattered among the cilia on the surface of a cell are trichocysts that shoot out of their retracted positions when stimulated, and they can be used for defense, procuring food, or for anchoring the cell (Figure 30.22).
e. Paramecium commonly represents the ciliates (Figure 30.22).
1. Paramecium has two kinds of nuclei: one or more small micronuclei provide genetic continuity from generation to generation and a large macronucleus, which provides the templates for protein synthesis and cellular regulation between divisions.
2. While Paramecium and other ciliates generally reproduce by asexual cell division, they occasionally engage in conjugation, or sexual mating (Figure 30.23).
30.7 The slime molds include a variety of amoeboid saprobes.
a. Slime molds are named for their glistening, slimy appearance (Figure 30.24).
1. Some species of slime molds are white, but most are red, orange, or yellow.
2. All slime molds feed and grow as amoeboid cells and then develop fruiting bodies that produce spores.
b. A true slime mold (phylum Myxomycota) grows vegetatively as a plasmodium, a large coenocytic mass that moves along slowly like a gigantic amoeba, phagocytizing organic particles as it goes.
1. The diploid plasmodium stops moving and sends up fruiting bodies in which masses of haploid spores form by meiosis (Figure 30.24).
2. After falling to the ground, these spores germinate into flagellated gametes, which fuse in pairs to make new diploid vegetative cells.
3. Each cell then begins to move about as an amoeba again.
c. Cellular slime molds (phylum Acrasiomycota) are unrelated to, but share a similar life cycle with the true slime molds.
1. The well-studied Dictyostelium consists of independent, uninucleate amoeboid cells that feed phagocytically as they move over the soil and divide repeatedly into more uninucleate cells (Figure 30.25).
2. The cells recruit one another and stream into a central point where they form a pseudoplasmodium, so called because the cells retain their cellular identities and do not form a coenocyte.
3. There are no sexual events in this cycle, and all stages are apparently haploid.
C. EVOLUTION AMONG THE BASAL EUCARYOTES
30.8 Eucaryotic cells and the mitotic apparatus arose through a series of complicated changes.
a. One hypothesis is that the eucaryotic condition was a response to an oxidizing atmosphere, created by phototrophs that produce oxygen as a byproduct.
b. A second hypothesis is that the eucaryotic cell, with a mitotic apparatus, evolved as housing for a large genome and as a mechanism for regularly dividing such a genome among daughter cells.
c. Because mitochondria and chloroplasts provide efficient mechanisms for obtaining energy, more complex, energy-demanding processes are possible.
d. Figure 30.26 shows a likely scenario for the evolution of a procaryotic cell into a eucaryotic cell.
e. Microtubules must have evolved early in the eucaryotic lineage.
30.9 Dinoflagellates and euglenoids have primitive types of nuclear structure and mitosis.
a. The discovery of primitive nuclear features and modes of mitosis among certain eucaryotes happened during the 1960s, when several investigators found the unusually primitive nuclear features of dinoflagellates.
1. John Dodge showed that the chromosomes of dinoflagellates look more like the nuclear bodies of bacteria than the chromosomes of other eucaryotes (Figure 30.27).
2. Donna Kubai and Hans Ris showed that some dinoflagellates undergo mitosis as shown in Figure 30.28, where the nuclear envelope does not break down.
3. The several chromosomes divide rather like those of bacteria, but with the aid of a new eucaryotic feature, microtubules.
b. Although mitosis in other basal eucaryotes is quite varied, the process has apparently evolved through two kinds of changes (Figure 30.29).
1. Kinetochores were initially plaques in the nuclear envelope to which the chromosomes were bound. Later the kinetochores became permanently attached to the chromosomes, and the nuclear envelope no longer had any role in mitosis.
2. The spindle was initially extranuclear, then some spindle elements began to separate the chromosomes by attaching directly to the kinetochores.
c. Primitive nuclei and mitosis also characterize the euglenoid flagellates (Figure 30.30).
1. Although their chromosomes are like those of other eucaryotes, their nuclei still divide by elongating and pinching in two, as the nuclei of dinoflagellates do.
2. They use the AAA pathway of lysine synthesis.
3. They have flagella with only a single row of filaments.
4. They have a gullet into which they take small bits of food.
5. They have a brightly colored eyespot near the gullet that is apparently a light receptor.
6. They have a peculiar squirming motion in which the cell alternately bulges and narrows.
d. The euglenoids have another peculiar feature - a rod running through each flagellum parallel to the microtubules.
1. The euglenoids are thought to be close relatives of the trypanosomes, a group that includes such parasites as the causative agent of African sleeping sickness (Figure 30.31).
2. The tsetse fly is a vector that carries these trypanosomes between humans and various animals.
30.10 Some eucaryotic organelles probably arose through endosymbiosis.
a. The endosymbiotic model theorizes that eucaryotic cells acquired mitochondria and chloroplasts by incorporating certain bacteria as endosymbionts, organisms that grow inside other cells (Figure 30.32).
1. This theory is mainly championed by Lynn Margulis.
2. According to this model, the primitive eucaryotic cell was fundamentally anaerobic and lived by fermentation.
3. Then some cells acquired endosymbionts that transformed their metabolism.
b. Although the endosymbiotic model is very plausible, some biologists favor an alternative model (Figure 30.32).
1. This model supports the idea that mitochondria could be derived from internal changes in a cell.
30.11 Can we derive a composite picture of early eucaryotic evolution?
a. Figure 30.33 summarizes several lineages of evolution among the protists and the other eucaryotic kingdoms.
b. According to the endosymbiotic model, various organelles were incorporated at quite different times, and events must have occurred quite independently in different clades (Figure 30.34).
c. A body of fascinating evidence indicates that red algae and fungi are related.
d. In 1993, Sandra Baldauf and Jeffrey Palmer presented evidence for a close relationship between fungi and animals.
e A group of simple fungi called chytrids present a puzzle for scientists.
1. Figure 30.35 shows the life cycle of the chytrid Allomyces.
2. Chytrids are not true fungi because they have flagella and they reproduce with motile spores powered by a single posterior flagellum.
f. Figure 30.36 shows one possible phylogeny of some important groups of organisms discussed in this chapter.
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