Sponges

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Phylum Porifera

-- 5000 -10000 spp.


-- mostly marine some fw
none terrestrial
-- 3 classes, most important
distinction are skeletal
characteristics
Phylum Porifera
I. General Ecological Characteristics
II. Body Plan
III. Metazoan Characteristics
A. Cell types
B. Allorecognition
C. Reproduction and Embryology
D. Other metazoan homologies.
IV. Sponge Phylogeny
I. General Ecological Characteristics
Sponges are:
• Sessile, benthic
• Filter feeders
• Competitors for space
• Fed upon by specialist predators
• Grow in many forms, solitary,
colonial, branching, as thin
sheets over substrates
• From few cm to over 1 m in size
• Estimated in some cases to be
several hundred years old
II. General Characteristics of
the Porifera Body Plan
The classification of sponges is based on skeletal morphology
images.slideplayer.com
II. General Characteristics of
the Poriferan Body Plan

3 major types of body construction


Asconoid
Syconoid
Leuconoid

(this has little to do with the


classification of sponges, which is
based on skeletal morphology)
Asconoid Sponge Syconoid Sponge
Osculum

Ostia
Leuconoid Arrangement
These are the largest, and most sponges
have this type of construction
Sponge Feeding Movie

Simulations of
Sponge Feeding

http://www.biology.ualberta.ca/facilities/multimedia/uploads/zoology/Porifera.swf

http://www.youtube.com/watch?
v=RmPTM965-1c&feature=related
Carnivorous
sponges from deep
water and shallow
caves

Evolution of
macrophagy from a
microphagous,
filter feeding life
style
II. Other Characteristics of
the Poriferan Body Plan
• No true muscular system
• Lacking sensory organs, nervous system
• Often amorphous and asymmetrical,
no anterior, posterior, oral surfaces
• No true tissues
• All physiological functions carried out \
at the cellular level
Begs the question: Colony of protista or
a simple metazoan (i.e. an integrated animal ?)
III. Metazoan-like Characteristics of Sponges
A. 5 different principal cell types (20 total)

Osculum
Pinacocyte

Choanocyte
spongocoel

Porocyte Sclerocytes

Archeocyte

http://www.biology.ualberta.ca/facilities/multimedia/uploads/zoology/Porifera.swf
Stem cells – have the capacity of self-replication
and to give rise to more than one type of
mature daughter cells

Image courtesy of BioMEDIA ASSOCIATES

Archeocytes – in sponge embryos are considered


totipotent stem cells that can give rise to an
entire organism
- in adults they produce a few cell types
(sclerocytes, germ cells, etc.) but not an
entire organism; they are considered
pluripotent
Muller (2006) Seminars in Cell and Developmental Biology #17:481-491
III. Metazoan-like Characteristics of Sponges
B. Complex reproduction
III. Metazoan-like Characteristics of Sponges
Unique but complex embryonic development, with a hollow
blastula stage but does not form a gastrula

Carried in the plankton


Settlement &
sperm metamorphosis
Parenchymula or
Amphiblastula

Egg Juvenile
Adult

Sexual reproduction involves fertilization, release of a planktonic


larva, and its eventual settlement and metamorphosis on the bottom.
III. Metazoan-like Characteristics of Sponges
C. Other sponge metazoan homologies: Epithelium

-- pinacoderm: epithelial-like layer homologous to animal


epithelia with collagenous sublayer ( ...but basal lamina
only in Homoscleromorphs)…

-- septate junctions (desmosomes) albeit primitive and small


(only Calcarea has full “animal-like” desmosomes)
-- extracellular matrix

-- spongin is collagen-like molecule

-- ubiquitin protein similarity (tag other proteins


for proteolysis)
Integrative and Comparative Biology Advance Access published online on June
22, 2009
Integrative and Comparative Biology, doi:10.1093/icb/icp038
Epithelia and integration in sponges

Sally P. Leys1,*, Scott A. Nichols{dagger} and Emily D. M. Adams*


*Department of Biological Sciences, University of Alberta, Edmonton, AB,
Canada T6G 2E9; {dagger}Department of Molecular and Cell Biology, University
of California, Berkeley, CA 94720, USA

Correspondence: 1E-mail: [email protected]

An epithelium is important for integrity, homeostasis, communication and co-ordination, and its
development must have been a fundamental step in the evolution of modern metazoan body plans. Sponges
are metazoans that are often said to lack a true epithelium. We assess the properties of epithelia, and
review the history of studies on sponge epithelia, focusing on their homology to bilaterian epithelia, their
ultrastructure, and on their ability to seal. Electron micrographs show that adherens-type junctions are
present in sponges but they can appear much slighter than equivalent junctions in other metazoans. Fine
septae are seen in junctions of all sponge groups, but distinct septate junctions are only known from
Calcarea. Similarly, all sponges can have collagenous sheets underlying their epithelia, but only
homoscleromorphs are established to have a distinct basal lamina. The presence of most, but not all, gene
families known to be involved in epithelial development and function also suggests that sponge epithelia
function like, and are homologous to, bilaterian epithelia. However, physiological evidence that sponge
epithelia regulate their internal environment is so far lacking. Given that up to six differentiated epithelia
can be recognized in sponges, distinct physiological roles are expected. Recognition that sponges have
epithelia challenges the perception that sponges are only loose associations of cells, and helps to relate the
biology and physiology of the body plan of the adult sponge to the biology of other metazoans.
III. Metazoan-like Characteristics of Sponges
C. Other sponge metazoan homologies:
Regulation of Development

-- True Hox genes are not found, but many homologous


developmental transcription factors are conserved
-- Most of the developmental signaling pathways
(Wnt, Notch) and they are expressed along the same
embryonic “axis” in sponges (and Cnidaria).
Many of these signaling pathways and transcription
factors have not been found in Protists.

From Adamaska et al., 2011


III. Metazoan-like Characteristics of Sponges
D. Non-self recognition
“Allorecognition”

Some sponges form new individuals hours after their


cells are separated from one another. If species are
combined, the cells segregate with their own
“Metazoan-like Characteristics of Sponges”
Allorecognition Histoincompatability

Immune response:
antagonism toward foreign substances
antagonism must be specific toward that substance
future responses should be altered by the first response

The sponge immune response is mediated by molecules which have


been found to control histo-recognition in deuterostomes
including Immunoglobulin-like domains and citokines
Summary
-- Sponges lack complexity, but their body plan is
ecologically and evolutionarily successful

--They should be considered metazoans


since they have fundamental characteristics of
multicellular animals;

--They are derived from flagellated protists but may be a


early and now distant branch of the metazoa;
animals are monophyletic
Phylum Placozoa
-- 2-3 mm, 25 um- thick, resembling a large ameba
-- Lacks anterior posterior polarity
-- Asexual reproduction is prevalent
--The most primitive animal?

Trichoplax adhaerens
Phylum Placozoa Dorsal cilium
epithelium-like layer

Fiber synctium

thick glandular layer


Intercellular
junctions
Flagellated cells
-- Feed ventrally by absorption of digested material
-- Lack organs but tissue-like outer walls (no basement membrane)
-- A bit more than 2000 cells
-- Only 4 different cell types (20 in sponges; > 220 in mammals)
-- Smallest genome of all animals
Three competing Scenarios

A. Earliest view of them as the basal metazoan


B. Special cellular junctions consisting of two opposing
dense plaques (desmosomes) not found in most sponges
C. 16S rRNA data…maybe secondarily simplified from more
complex ancestors?
Mitochondrial genome of
Trichoplax adhaerens
supports Placozoa as the
basal lower metazoan phylum
Dellaporta, Stephen L.
et al. (2006) Proc. Natl.
Acad. Sci. USA 103,
8751-8756

Largest known mt genome 46


kbp, 2x that of most metazoa
with introns and other
intrageneic spacers and large
protein coding regions that are
usually lacking in other animals

Blue: known mt proteins


Gray: ribosomal genes
Green: unknown open reading frames
Red lines: introns
(a sponge)

Conclusions?

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