Protist Notes

Protists

Overview

  • Protists are mostly single-celled eukaryotes. They are not plants, animals, or fungi.
  • This group is not a kingdom because some protists are more closely related to plants, fungi, or animals than other protists.
  • Protists and other eukaryotes possess cells with a nucleus and other membrane-enclosed organelles.
  • Protists account for a significant portion of eukaryotic diversity.
  • Most organisms in eukaryotic lineages are protists.
  • The majority of protists are unicellular.

Nutritional Diversity

  • Protists exhibit the most nutritional diversity among eukaryotes.
    • Photoautotrophs contain chloroplasts.
    • Heterotrophs absorb organic molecules or ingest larger food particles.
    • Mixotrophs combine photosynthesis and heterotrophic nutrition.

Modes of Reproduction

  • Some protists reproduce asexually only.
  • Others have both asexual and sexual phases in their life cycle.
  • All three basic types of sexual life cycles (animal, plant, and fungal) are represented among protists.

Alternation of Generations

  • Multicellular algae exhibit a variety of life cycles.
  • Some have alternation of generations, where both haploid and diploid stages are multicellular.
  • The diploid generation is called a sporophyte because it produces spores.
  • Haploid spores develop into multicellular haploid gametophytes that produce haploid gametes.
    • Heteromorphic species (e.g., Laminaria) have structurally different gametophytes and sporophytes.
    • Isomorphic species have gametophytes and sporophytes that look similar.
  • Fertilization of gametes results in a diploid zygote, which develops into a new sporophyte.

Laminaria Life Cycle

  • Haploid (n) and Diploid (2n) stages alternate.
  • Sporophyte (2n) produces sporangia via meiosis.
  • Sporangia release zoospores.
  • Zoospores develop into male and female gametophytes (n).
  • Gametophytes produce sperm and eggs.
  • Fertilization results in a zygote (2n).
  • Zygote develops into a new sporophyte.

Endosymbiosis

  • Endosymbiosis is a symbiotic (mutualistic) relationship in which one organism lives within another.
  • The first eukaryotes were likely heterotrophs.
  • All eukaryotes have mitochondria, but not all have plastids (plant organelles).
  • Serial Endosymbiosis: A eukaryotic heterotroph absorbs a proteobacteria, which becomes mitochondria, then absorbs a photosynthetic cyanobacteria, resulting in a eukaryote that can now photosynthesize.
    • Examples: euglena, brown algae, diatoms, red algae, green algae, plants.

Evidence for Endosymbiosis

  • Mitochondria and plastids have their own DNA.
  • They can transcribe and translate their own DNA into proteins, using a slightly different genetic code than the eukaryotic host.
  • Inner membranes of mitochondria and plastids have enzymes and transport systems homologous to those in bacteria.
  • Replication of mitochondria and plastids occurs via binary fission.
  • The size, RNA sequences, and antibiotic sensitivity of mitochondrial/chloroplast ribosomes are more similar to bacteria than eukaryotic cytoplasmic ribosomes.

Secondary Endosymbiosis

  • Red and green algae were ingested by heterotrophic eukaryotes, a process called secondary endosymbiosis.
  • "Algae" is not a valid taxonomic name.
  • The following can all be considered "algae": Protist, Algae, Plankton

Supergroups Including Algae

  • SAR (Stramenopiles, Alveolates, Rhizarians)
  • Excavata: Euglenozoans
  • Rhizara: Cercozoa

Colonies vs. Multicellularity

  • Colonies: Collections of cells connected to one another, but all cells are roughly the same; this is NOT true multicellularity.
  • Colonial species are often found in fossils.
  • Living colonial species usually have rigid cell walls shared to form the colonies.
  • Colonies are also found in prokaryotes.

Examples of Colonies

  • Volvox

True Multicellularity

  • True multicellularity involves differentiated cells.
    • Examples: Red, green, and brown algae, plants, fungi, animals.
  • Complex multicellularity has evolved multiple times in evolutionary history.
    • Animals have a single-celled common ancestor.
    • Fungi have a different single-celled common ancestor.
    • Thus, animals and fungi evolved multicellularity independently.
  • Plants, Animals, and Fungi are truly multicellular.
  • Some algae (e.g., brown algae, green algae) independently derived multicellularity, different from that in plants.

Eukaryotic Diversity

  • 'Protists' is not a taxonomically valid group.
  • They have not been a monophyletic clade since the 1960s.
  • "Kingdom Protista does not exist."
  • 'Protists' are probably nonmonophyletic because of multiple independent events of endosymbiosis.
  • Different protist lineages are actually the ancestral forms of multicellular eukaryote kingdoms: plants, animals, and fungi.

Major Supergroups of Eukaryotes

  • Phylogeny is supported by genetic systematics.
  • There is significant morphological diversity within supergroups, making it challenging to align morphological traits with molecular phylogenies.
  • It is difficult to tell an evolutionary story based solely on morphology.
  • These protist supergroups do NOT share a common ancestor.

Supergroup Excavata

  • General Feature: "Excavated" groove on one side of the cell body.
  • Clades:
    • Parabasilids and Diplomonads have highly reduced mitochondria (adapted for anaerobic conditions).
    • Euglenozoans have unique flagella.
      • Examples: Trichonympha, Trypanosoma, Euglena.

Excavata: Diplomonads and Parabasilids

  • General Features:
    • Reduced mitochondria that do NOT have electron transport chains.
    • Cannot use oxygen; must use anaerobic respiration.
    • Many are parasites.
  • Examples:
    • Diplomonad: Giardia (intestinal parasite that causes Giardiasis).
    • Parabasilid: Termite gut mutualist Trychonympha.

Excavata: Euglenozoans

  • General Features:
    • Unifying morphological feature: Flagella composed of an internal supporting rod.
    • Extremely diverse; includes heterotrophs, autotrophs, parasites.
  • Subclades:
    • Kinetoplastids have a kinetoplastid (giant mitochondria).
      • Aquatic heterotrophs and parasites.
      • Example: Trypanosoma (causes African Sleeping Sickness, via Tsetse fly as vector).
    • Euglenids have a 'pocket' at one end where 1 or 2 flagella emerge.
      • Euglena can switch between photosynthesis and heterotrophy.

Supergroup SAR

  • SAR is a monophyletic supergroup named for the first letters of its three major clades: Stramenopiles, Alveolates, and Rhizarians.

SAR: Stramenopiles (“Straw Hair”)

  • General Features: Straw-like hair projections found in the long flagellum.
  • Ecology and Evolution:
    • Stramenopiles share a common ancestor with Alveolates.
    • Some photosynthesizers in this group possibly gained the ability to carry out photosynthesis via secondary endosymbiosis of a heterotrophic protist engulfing a red algae.
  • Stramenopiles includes: Diatoms and Brown Algae.

SAR: Stramenopiles: Diatoms

  • Anatomy:
    • Unique perforated silicon “armor” cell walls, which have two parts that overlap like the two halves of a Petri dish.
  • Ecology:
    • Diatoms are diverse and abundant unicellular phytoplankton; they are photosynthetic protists.
    • With over 100,000 species – possibly one of the most important photosynthesizers on Earth.
  • Human Application:
    • "Diatomaceous earth" is fossilized remains of diatoms.
    • Applications include filtration aid, mild abrasive (e.g., toothpaste), absorbent for liquids, reinforcing filler in plastics and rubber, anti-block in plastic films, cat litter, activator in blood clotting studies, a stabilizing component of dynamite, a thermal insulator, soil for potted plants and trees (e.g., bonsai).

SAR: Stramenopiles: Brown Algae

  • Anatomy, Physiology, and Reproduction:
    • Largest and most complex algae; almost all species are multicellular.
    • All are marine, typically cold-water.
    • Can carry out alternation of generations.
    • Carotenoids in plastid are dominated by brown pigments named fucoxanthin.
    • Brown algae carry out photosynthesis on their blades.
    • Photosynthesis occurs via chlorophyll a and c (there is no chlorophyll b).
    • Excess glucose is stored as complex polysaccharides, known as laminaran (unlike plants which store glucose as starch).

SAR: Alveolata

  • General Features: Have membrane-enclosed sacs (alveoli) just under the plasma membrane.
  • Systematics: Includes 3 clades: Dinoflagellates, Ciliates, Apicomplexans.

SAR: Alveolata: Dinoflagellates

  • General Features:
    • Unicellular.
    • Reinforced cellulose “armor” plates with 2 flagella in the groove between plates.
    • Some dinoflagellates release toxins.
  • Locomotion: Peculiar rolling motion caused by the 2 flagella.
  • Ecology:
    • Includes autotrophs, heterotrophs, and mixotrophs.
    • Toxins released by dinoflagellates are usually harmless at small population densities but dangerous at high population densities, which can kill fish and molluscs (and close beaches).
    • Huge accumulations can lead to red tides (appear red because of carotenoids).

SAR: Alveolata - Ciliates

  • General Feature: Use cilia to move and feed.
  • Two types of nuclei: macronucleus and micronucleus.
  • Anatomy and Ecology:
    • Unicellular.
    • Most are heterotrophic predators (e.g., Paramecium).
  • Reproduction:
    • Reproduce asexually via binary fission.
    • Genetic variation accomplished by conjugation.

SAR: Alveolata: Apicomplexans

  • General Feature: Almost all are animal parasites that possess a complex of organelles at one end (“apex”) of the cell that facilitates entry into a host cell.
  • Physiology: Parasitic; NOT photosynthetic.
  • Reproduction: Often require multiple hosts.
  • Malaria life cycle: Plasmodium vivax causes malaria. It travels via mosquitoes (vector) to the final host, humans.

SAR: Rhizarians

  • General Feature: Possess very thin pseudopodia.
  • Anatomy and Physiology:
    • Some are amoeboid that move and feed using these thin pseudopodia.
    • Others are flagellated non-amoeboids that use pseudopodia mainly for feeding.
  • Systematics: Major clades: Foraminifera (Forams) and Cercozoans.

SAR: Rhizarians: Foraminiferans (Forams)

  • General Feature: Have porous shells called tests (“armor” made of calcium carbonate).
  • Forams can extend their thin pseudopodia through holes.
  • Ecology:
    • Mostly marine, but also found in brackish, freshwater, and even terrestrial habitats.
    • Benthic or planktonic.
    • Some form symbioses with photosynthetic algae.

SAR: Rhizarians: Cercozoans

  • General Features:
    • First identified in molecular phylogenies.
    • Amoeboid or flagellated.
    • Thread-like pseudopodia for feeding.
  • Ecology:
    • Many free-living (many parasitic).
    • Marine, freshwater, soil.
    • Mixotrophs, heterotrophs (predators, parasites), autotrophic (photosynthetic).

Supergroup Archaeplastida

  • Includes red algae, green algae (2 lineages, one of which evolved into plants).
  • Most recent common ancestor of this supergroup was probably a heterotrophic protist that absorbed a cyanobacterium (which became the chloroplast).

Archaeplastida: Red Algae

  • General Features:
    • Red algae have red photosynthetic pigment phycoerythrin.
    • Store excess glucose in carbohydrate form outside of the chloroplast as starch.
  • Anatomy and Ecology:
    • Photosynthetic autotrophs.
    • Generally multicellular.
    • Chloroplasts contain chlorophyll a.
    • Store excess glucose in carbohydrate form outside of the chloroplast as starch.
    • Isomorphic alternation of generations.
  • Human Interest:
    • In addition to cellulose, their cell walls contain carrageenan, used as a thickener in foods, and agar, used as a microbiological growth medium.
    • Dried red algae is also wrapped around rice in sushi.

Archaeplastida: “Green Algae”: (Chlorophytes and Charophytes)

  • General Features:
    • Photopigments are chlorophyll a, chlorophyll b, and carotenoids.
    • Excess glucose is stored as amylose starch contained within the chloroplast.
    • Cell walls composed of cellulose.
    • Charophytes (NOT Chlorophytes) are the sister group to plants.
    • "Plants" are defined as being multicellular and can live on land (In contrast, algae always need to be in a watery/moist environment).
  • Green Algae:
    • "Green algae" is a paraphyletic grouping.
    • Unicellular, colonial, multicellular.
    • Have complex life cycles with sexual and asexual stages.
    • Chlorophytes have alternation of generations:
      • Ulva (a chlorophyte) has isomorphic alternation of generations (sporophyte and gametophyte generations look exactly alike).
      • Derbesia (a chlorophyte) has heteromorphic alternation of generations.
    • Strangely, charophytes do not have alternation of generations.

Supergroup Unikonta

  • Includes Animals, Fungi, and other eukaryotes.
  • Two major clades: Amoebozoans (tubulinids and slime molds) and Opisthokonts (animals, fungi, and closely related ‘protists’).

Unikonts: Amoebozoans

  • General Features:
    • Amoeba-like protists.
    • Compared to rhizarians, amoebozoans have thicker pseudopodia that can be lobe-shaped or tube-shaped.
    • Cytoplasmic streaming.
  • Ecology: Most are predaceous heterotrophs.
  • Systematics:
    • “True” amoeba belong to this group.
    • Includes slime molds: slime molds were once thought to be part of fungi, but are NOT fungi; slime molds have different unicellular ancestors than fungi.

Unikonts: Opisthokonts

  • Includes animals and their protist relative (choanoflagellates) and fungi and their protist relative (nucleariids).
  • Thus, the common ancestor of plants, fungi, and animals was probably a single-celled eukaryote, suggesting the independent evolution of multicellularity in all three multicellular lineages.
  • Conclusion: Multicellularity in plants, fungi, and animals resulted from convergent evolution.

Protists as the Base of Aquatic Food Chains

  • Many different photosynthetic ‘protists’ comprise the base of aquatic food chains.
    • Ciliates
    • Forams
    • Euglenozoans