SEM of Paramecium

Explain, in the context of evolution, why there is more

higher order diversity (e.g. phyla) in the ocean but a
greater species diversity on land.
Kingdom
Protista

(Based on Textbook Chapter 3)

Protista and the Origin of Metazoans

Protista Defy Tidy Definition:
Goals:
Review characteristics of single
cell body plan
Explore the limitations of life
as a single cell
Learn some of the ways in which
protists are important
Evaluate evidence for the origin
of animals from protist ancestors
Introduction - >10 phyla, believed by
I. Body Plan some that they should be
A. S.A./Volume Constraint
classified as Kingdoms
B. Reynolds number
- Among them are the
II. Important Protists
ancestors of multicell
A. In food webs organisms including
B. Agents of disease animals.
III. Origin of Metazoans
Evidence for colonial origin
and metazoan ancestors

Body plan characterized by: Most Common TYPES of Protista:

Ciliates (Phylum Ciliophora): externally ciliated
A. Unicellularity (most)
B. Small Size (< 5-250 µm) Sarcodinids: 4 phyla of amoeboid types
C. Lacking Organ Systems (Amoebas, foramenifera, radiozoans, heliozoans)
(including a nervous system)
Flagellates: several phyla of flagella-bearing
types, including many photosynthetic forms and
Each individual is a cell that is Some important parasitic forms
functionally more versatile than any
single cell from a multicell organism
Spore-Formers: Phylum Apicomplexa and other
very important parasitic forms

1
Means of locomotion and associated structures are no longer
considered key characters in establishing the phylongeny
Kingdom Protozoa
(evolutionary relatedness) of Protista. Phylogeny is Complicated/Unresolved

Alveolate Amoeboid Flagellated

Protozoa Protozoa Protozoa
Paramecium

Ciliates EUGLENOZOA
foramenifera
PH CILIOPHORA
dinoflagellate GRANULORETICULOSA trypanosomes
(ciliates) (foramenifera) euglinids
Euglenoid DINOZOA RADIOZOA trichomonads
(dinoflagellates) (radiolarians)
Giardia
APICOMPLEXA ACANTHARIA
(coccidians (actinarians)
e.g. Plasmodium) HELIOZOA
Didinium
Amoeboid Flagellated (heliozoans)

See pie chart ad taxonomic detail on pg 75

Protista and the Origin of Metazoans

I. Protistan Body Plan is Characterized by

Introduction -- Primarily unicellular

I. Body Plan -- Small, 5 -300 µm
A. S.A./Volume Constraint -- Lack organs
B. Reynolds number (Lack a nervous system)
II. Important Protists
A. In food webs
B. Agents of disease What constraints does the small unicellular body
plan impose on the biology of protistans?
III. Origin of Metazoans
Evidence for colonial origin
and metazoan ancestors

S.A./Vol relationship imposes a limit on cell size

As a cell increases in size,
exchange with the environment
Some Amebas
becomes more difficult
partly solve
1 this problem
Inch
L Surface Area ~ L2 (2.5 cm)
Volume ~ L3
Therefore S.A./V declines
L as L increases

Cytoplasmic streaming provides continuous

mixing or circulation of the cytosol

2
Small Size also affects scaling of physical forces Re = velocity x length of the organism
Viscocity of Medium
Balance of inertia and drag When Re is high, drag is relatively negligible
on a moving organism (example, Re is 30,000 for a dragonfly in flight)

Depends on the size and velocity of the organism When Re is low, drag forces dominate movement
and on the viscocity of the medium. (example, Re for a protist could be <0.1)
Expressed as Reynolds Number (Re)
Re = velocity x length of the organism In what way would low Reynolds
Viscocity of Medium numbers affect the ecology of
Re vel x L protists?
=
Vis

Re = velocity x length of the organism

Viscocity of Medium

What is life like at Low Reynolds numbers??

-- no inertia
-- force of gravity balanced by viscocity
-- yet ciliates travel 2 mm per sec. (8 body lengths)
-- feeding also affected

Vorticella feeding

Re = vel x L I. Protistan Body Plan is Characterized by

Vis
-- Primarily unicellular
-- Small, 5 -300 µm
Viscous forces can -- Lack organs
also be important in (Lack a nervous system)
air but the medium
is not very dense
what conditions will
What constraints does the small unicellular
produce a low Re???
body plan impose on the biology of protistans?

Midge

3
 Anterior
Organelles are the Food
functional equivalents of vacuole

organs.

Oral
nuclei groove
Organelles not generally
found in metazoans
Cyto-
- contractile vacuoles stome
- trichocysts
- toxicysts
Fig 3.19
contractile
Posterior Contraction
http://www.youtube.com/watch?v=hSFJ-
vacuole
Paramecium Feeding spnn6c&NR=1

II. Importance of Protozoa

Spasmin filaments

The spasmoneme is a biological
spring organelle. Contraction is
completed in 2 ms at speeds up
to 8 cm/sec and is triggered
by Ca++ activated interactions
along the length of the
supramolecular structure. The
reaction does not require ATP
A. Major role in ecosystems as decomposers, food source for
metazoans
B. Responsible for a large number of human ailments, including
malaria, dysentery, sleeping sickness
C. Provided biologists with outstanding material for genetic,
physiological and other studies.

Protista and the Origin of Metazoans

Metazoa arose from Protozoa sometime
Introduction
before the Cambrian 1200-600 mya
I. Body Plan
Which protozoan group was the ancestor??
A. S.A./Volume Constraint
B. Reynolds number
By what process was first metazoan formed??
II. Important Protists
A. In food webs Are metazoans Monophyletic or Polyphyletic??
B. Agents of disease

III. Origin of Metazoans What were the first Metazoans like??

Evidence for colonial origin
and metazoan ancestors CAN WE EVER REALLY KNOW??

4
 3 Phyla are Possible Candidates
as Basal Group of all Animals
What is a “metazoan”??

Eukaryotic and multicellular

Cell differentiation and coordination;

more than a colony or aggregation of cells
Cnidarians
Layers of cells organized into “tissues”
Sponges
Embryonic development; gastrulation

Flatworms

Symbiotic Hypothesis
Theoretically
3 ways in which
Colonial Hypothesis
a multicellular
organism could
evolve from a
protist.
Cellularization Hypothesis

Strengths: Origin of Eukaryotes from prokaryotes

Weakness: Genetic obstacles to reproduction. How
do they integrate into a reproducing organism?

Cellularization
Stained Paramecium showing multiple ”inclusions”

Macronucleus

Strengths: Paramecium-flatworms are similar

Weaknesses: no evidence of “compartmentalization”
:flatworms with complex embryology
:What about Cnidaria, Sponges ??

5
 Similarities of Choanocytes (sponge cells)
and choanoflagellates (protozoan)

(Haeckle’s favorite)

Blastea

Gastrea

Strengths: models exist (Volvox, choanoflag.)

: parallels animal embryology
Weaknesses: ??? Choanocyte Choanoflagellate

Model organization

Eudorina

Pandorina

Choanoflagellates live as solitary and colonial forms Volvox

Genetic Evidence (18S r-DNA)

Opisthokonts share:
uniflagellate (reproductive) stages with flagellum
at the posterior
Mitochondria with flattened, plate-like cristae

6
Based on sequence
analyses of 5 proteins
Monosiga has 86%
sequence homology with
animals.

a. Phylogeny based on protein

domain comparisons

b-d. choanoflagellate cells bear a single

Flagellum and an apical collar of actin-
filled microvilli (bracket, c). D. an
overlay of b-tubulin (reen), polymerized
Mol. Biol. Evol. actin (red) and DNA (blue) localization
23: 93-106 2006 reveals the position of the flagellum
within the collar of microvilli.

Conclusions on Metazoan Origin

Evidence best supports the colonial theory

Choanoflagellate group is most likely ancestor;

Evidence includes cell structure and genetic
similarities

Idea that metazoans are monophyletic is supported