Major Evolutionary Transitions and Lifestyles

How many times did multicelluarity appear to evolve?

a. once
b. twice
c. many times
How many times did multicelluarity appear to evolve?

a. once
b. twice
c. many times
Was the transition to multicelluarity evolved several ?
General features of multicellular origins in evolution:

Aggregative multicellularity

Clonal multicellularity
 Red Algae
A Cryptophytes Glaucophytes
Cyanidiophytes
Katablepharids Bangiophytes

**
Picozoa Porphyridiophytes
Palpitomonas Floridiophytes
Centrohelids
Telonemids Trebouxiophytes
Haptophytes Chlorophyceans
Rappemonads Ulvophytes Viridiplantae
Dinoflagellates Prasinophytes
Syndiniales Mesostigma
Alveolates Oxyrrhis Zygnemophyceans
Perkinsus Charophyceans
Land Plants
Apicomplexa
Vitrella
Colpodellids
Bryophytes
Tracheophytes * *
Kinetoplastids
Chromera Diplonemids
Colponemids Euglenids
Ciliates Heteroloboseans
Bicosoecids Jakobids
Labyrinthulids Trimastix Excavates
Thraustochytrids Oxymonads
Blastocystis Parabasalids
Actinophryids Retortamonads
Oomycetes Diplomonads
Bolidophytes
Diatoms Malawimonas
Stramenopiles
* Phaeophytes
Dictyochophytes
Pelagophytes
Collodictyon
Ascomycetes
Basidiomycetes **
*
Eustigmatophytes Zygomycetes Fungi
Pinguiophytes Chytrids
Raphidophytes Microsporidia
Chrysophytes Cryptomycota
Synurophytes Fonticula
Types of Xanthophytes Nucleariids Opisthokonts
multicellularity Chlorarachniophytes Ichthyosporea
Guttulinopsis
Clonal multicellularity Cercomonads Filasterea
Euglyphids
Phytomyxea Choanoflagellates
Aggregative multicellularity Gromia

Rhizaria
Haplosporidia
Mikrocytos
Acantharea
Metazoa
Breviata
Apusomonads
* Holozoa

Clade contains both Polycystines Ancyromonads

unicellular and multicellular Foraminifera Arcellinids
Archamoebae Tubulinids
species Myxogastria Leptomyxids
Dictyostelia Vannellids
* “Complex multicellularity” Amoebozoa

B Aggregative forms C Clonal forms

Evolution is not linear!

All different forms of multicellularity has

evolved at least 16 times!
Evolutionary Invention
Sponges: The evolution of multicellularity in animals
remains a mystery
 Solitary Choanoflagellate Colonial Choanoflagellate

Flagella
Hetrotrophs
Draw in food through water currents

http://www.cytographics.com/cg.html#
 Choanoflagellate-like
Solitary Choanoflagellate structures in the sponges

Flagella
Hetrotrophs
Draw in food through water currents
Nicole King
 Grassé, 1973) and in
scopy study of the de
E Choanocyte F (Weissenfels, 1982), t
far in transmission e
vane nofluorescence studi
vane basal foot here.) mt, mitochondr
(F and G) Basal mic
microvilli flagellum in choano
following Garrone (1
(1995), and Leadbeat
F G
1 μm

Golgi and the collar cell

G
Choanocyte
of some cnidarian
nucleus mt
basal foot cles produced b
nucleus (Goldberg and Ta
fv mt
and ctenophore
fv through endocyto
terocytes lining th
display a motile
(packed into a de
microtubules transition zone than forming a r
centriole 1991; Takashima
with a possible d
Brunet and King. 2017. The Origin of Animal Multicellularity and Cell Differentiation.collar cells.
Developmental Cell. DOI:https://doi.org/10.1016/j.devcel.2017.09.016
t the collar is a choanozoan synapomorphy. Interest- Finally, the homology of the collar
 (E) Comparative ultrastructural schematics of a
choanoflagellate and a sponge choanocyte, modi-
fied from Maldonado (2004) following Woollacott
fv and Pinto (1995) for the microtubule cytoskeleton
and Karpov and Leadbeater (1998) for the actin
cytoskeleton. (Although filopodia may occasionally
be present in choanocytes, as reported in sketches
from earlier studies of calcareous sponges (notably
2 μm Sycon raphanus) (Duboscq and Tuzet, 1939;
Grassé, 1973) and in one scanning electron micro-
scopy study of the demosponge Ephydatia fluviatilis
E Choanocyte
Tyrosine kinases
F (Weissenfels, 1982), they have not been reported so
far in transmission electron microscopy or immu-
vane nofluorescence studies and are thus not indicated
vane basal foot here.) mt, mitochondria.
(F and G) Basal microtubular foot supporting the
microvilli flagellum in choanoflagellates and choanocytes,
following Garrone (1969), Woollacott and Pinto
(1995), and Leadbeater (2014).
F G
Cadherins
1 μm

Golgi and the collar cells lining the gastrodermis

G
Choanocyte
of some cnidarians endocytose food parti-
nucleus mt cles produced by extracellular digestion

Integrins
basal foot
nucleus (Goldberg and Taylor, 1989). In bilaterians
fv mt
and ctenophores, nutrient acquisition
fv through endocytosis is performed by en-
terocytes lining the midgut that frequently
display a motile flagellum and microvilli

Extracellular Matrix
(packed into a dense brush border rather
microtubules transition zone than forming a ring; Hernandez-Nicaise,
centriole 1991; Takashima et al., 2013), consistent

e collar is a choanozoan synapomorphy. Interest-

Domains
with a possible derivation from ancestral
collar cells.
Finally, the homology of the collar complex in animals and
tozoan Ministeria vibrans (belonging to Filastera, choanoflagellates is supported by its restriction to choanozoans.
up of Choanozoa) sports microvilli-like tentacles Although morphologically analogous forms exist in a few
over its entire cell cortex (Cavalier-Smith and distantly related species (Mah et al., 2014), in all cases the under-
suggesting that microvilli might be more ancient lying ultrastructure differs. For example, in the amoebozoan
r complex. Phalansterium (Cavalier-Smith et al., 2004), the flagellum is sur-
Brunet and King. 2017. The Origin of Animal Multicellularity and Cell Differentiation.
conserved ultrastructure, the idea that the collar rounded by a ‘‘collar’’ formed by a continuous fold of cytoplasm
ved in choanozoan Developmental Cell.
ancestors is further DOI:https://doi.org/10.1016/j.devcel.2017.09.016
supported rather than by independent microvilli (Hibberd, 1983). In the ped-
stribution in animals. Beyond sponges, collar com- inellales Actinomonas and Pteridomonas (stramenopiles from
Coprinus Mushroom
 Development of the small mushroom Coprinus:

Agar medium

Hyphae grow

Spores germinate

Mushroom

Hyphae
anastamose
Sea Lettuce Ulva
Sea Lettuce Ulva
Green Alga - Volvox
Green Alga - Volvox
 Colonial Diatom

single individual

colony
Colonial Diatom
Evolutionary Invention
Zoothamnium: a colonial ciliate
Zoothamnium: a colonial ciliate
Evolutionary Invention
JOHN TYLER BONNER
1920 – 2019
 Cellular Slime Mold - Dictyostelium discoideum

Individual Amoebae feeding Food runs out - aggregate Fruiting bodies - spores

http://www.cytographics.com/cg.html#
Please watch and consider sharing this timeless
video, which I believe is the same one John
showed to Albert Einstein.
Laura Katz (Smith College, 09 February 2019)
 Red Algae
A Cryptophytes Glaucophytes
Cyanidiophytes
Katablepharids Bangiophytes

**
Picozoa Porphyridiophytes
Palpitomonas Floridiophytes
Centrohelids
Telonemids Trebouxiophytes
Haptophytes Chlorophyceans
Rappemonads Ulvophytes Viridiplantae
Dinoflagellates Prasinophytes
Syndiniales Mesostigma
Alveolates Oxyrrhis Zygnemophyceans
Perkinsus Charophyceans
Land Plants
Apicomplexa
Vitrella
Colpodellids
Bryophytes
Tracheophytes * *
Kinetoplastids
Chromera Diplonemids
Colponemids Euglenids
Ciliates Heteroloboseans
Bicosoecids Jakobids
Labyrinthulids Trimastix Excavates
Thraustochytrids Oxymonads
Blastocystis Parabasalids
Actinophryids Retortamonads
Oomycetes Diplomonads
Bolidophytes
Diatoms Malawimonas
Stramenopiles
* Phaeophytes
Dictyochophytes
Pelagophytes
Collodictyon
Ascomycetes
Basidiomycetes **
*
Eustigmatophytes Zygomycetes Fungi
Pinguiophytes Chytrids
Raphidophytes Microsporidia
Chrysophytes Cryptomycota
Synurophytes Fonticula
Types of Xanthophytes Nucleariids Opisthokonts
multicellularity Chlorarachniophytes Ichthyosporea
Guttulinopsis
Clonal multicellularity Cercomonads Filasterea
Euglyphids
Phytomyxea Choanoflagellates
Aggregative multicellularity Gromia

Rhizaria
Haplosporidia
Mikrocytos
Acantharea
Metazoa
Breviata
Apusomonads
* Holozoa

Clade contains both Polycystines Ancyromonads

unicellular and multicellular Foraminifera Arcellinids
Archamoebae Tubulinids
species Myxogastria Leptomyxids
Dictyostelia Vannellids
* “Complex multicellularity” Amoebozoa

B Aggregative forms C Clonal forms

Stuart A. West et al. PNAS
2015
 Why multicellularity?

What are the advantages?

Potential advantages of multicellularity remain unknown:

• resistance to environmental stressors with the outer cells shielding the inner
cells from harmful agents such as UV and toxic chemicals

• in terrestrial species, formation of a stalk allowing elevation from the substrate

and wind dispersal of the propagules.

• resistance to predators

• cooperative feeding

• division of labor (e.g., between flagellated cells and dividing cells)

• formation of a milieu interieur (i.e., a controlled ‘‘internal environment’’ of

stable composition).
What happens when mutlicelluarity breaks down?
Cancer is a highly
conserved feature
of animals

Abrams et al. 2009

Cancer is a highly
conserved feature
of animals

p53

cancer

Abrams et al. 2009