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Porifera (Sponges): Recent Knowledge and New Perspectives

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DOI: 10.1002/9780470015902.a0001582.pub2

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Porifera (Sponges): Introductory article

Recent Knowledge and . Introduction

Article Contents

New Perspectives . Diversity and Phylogeny

. Ecology

. General Organisation of the Body and the Aquiferous

Emmanuelle Renard, Institut Méditerranéen de Biodiversité et d’Ecologie marine et System
continentale (IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France . Cell Plasticity, Stem Cells and Cell Renewal

Eve Gazave, Institut Méditerranéen de Biodiversité et d’Ecologie marine et continentale . Reproduction and Development
. New Insights and Perspectives on Early Animal
(IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France Evolution

Laura Fierro-Constain, Institut Méditerranéen de Biodiversité et d’Ecologie marine et

Online posting date: 9th December 2013
continentale (IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France
Quentin Schenkelaars, Institut Méditerranéen de Biodiversité et d’Ecologie marine et
continentale (IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France

Alexander Ereskovsky, Institut Méditerranéen de Biodiversité et d’Ecologie marine et

continentale (IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France
Jean Vacelet, Institut Méditerranéen de Biodiversité et d’Ecologie marine et continentale
(IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France
Carole Borchiellini, Institut Méditerranéen de Biodiversité et d’Ecologie marine et
continentale (IMBE), UMR CNRS IRD Avignon Université, Aix-Marseille Université, Marseille, France

Based in part on the previous version of this eLS article ‘Porifera’ (2001)
by Patricia R Bergquist.

Porifera, commonly named sponges, are true animals or
metazoan despite their anatomical and morphological
simplicity that had led to a long-time debate about their
nature (animal versus vegetal) and constitution (colonies
of unicellular organisms versus multicellular organisms).
Sponges are a successful group of mostly marine filter
feeder organisms that represent a major life form of sev-
eral ecosystems. Devoid of organs, the main character-
istics of their body plan are the presence of a network of
pores (at the origin of their name), choanocyte chambers
and canals devoted to water filtration and spicules (when
present) allowing tissue physical support. Currently con-
sidered as the sister group to all the other animals, these
organisms have a key phylogenetic position. Recent stu-
dies have shown that they possess an unexpected
eLS subject area: Evolution & Diversity of Life
How to cite:
Renard, Emmanuelle; Gazave, Eve; Fierro-Constain, Laura;
Schenkelaars, Quentin; Ereskovsky, Alexander; Vacelet, Jean; and
Borchiellini, Carole (December 2013) Porifera (Sponges): Recent
Knowledge and New Perspectives. In: eLS. John Wiley & Sons, Ltd:
Chichester.
DOI: 10.1002/9780470015902.a0001582.pub2
molecular complexity raising exciting questions about
early animal evolution.
Introduction
Sponges or Porifera were already described by Aristotle,
and for almost 4000 years they have been used for human
bathing and cleaning purposes. As adults, sponges are
sedentary filter feeder animals that have considerable
ecological, commercial and biopharmaceutical value. They
can be found in all marine habitats and some of them
colonised freshwater. Filtration is performed by highly
specialised flagellated cells: the choanocytes, which enable
to pump the water unidirectionally through the body. The
water flow is canalised in an aquiferous system composed
of pores (ostia), canals and choanocyte chambers. The
aquiferous system not only ensures nutritive functions
but is also involved in other vital physiological functions,
such as reproduction and excretion. Sponges are devoid of
digestive cavity and gonads, and despite the lack of neu-
rons and muscles, they have the capacity to react to
environmental changes. Because of this anatomical sim-
plicity, they have been set apart from other metazoans for a
long time. They have also been considered as primitive
animals with a cellular instead of tissular organisation
level unlike other animals. Recent phylogenetic, cellular,

eLS & 2013, John Wiley & Sons, Ltd. www.els.net 1

 Porifera (Sponges): Recent Knowledge and New Perspectives
molecular and developmental data tend to propose a
re-evaluation of these classical interpretations.
Diversity and Phylogeny
Sponges are known and used by humans since the antiquity
(Pronzato and Manconi, 2008). According to the fossil
record, Porifera were already present and diversified in the
lower Cambrian (Rigby and Hou, 1995). In present time,
sponges have colonised all marine habitats and ecosystems,
and some species even managed to colonise freshwater.
Their diversity in terms of species number is quite difficult
to estimate precisely, notably not only because of the
scarcity and the variability of morphological characters
they harbour but also due to the difficulties to access their
diverse habitats until recently. As a result, recognition and
description of new species is a highly specialist task. In this
context, molecular data, in addition to the more classic
morphological, embryological, and cytological data, have
been shown to be an invaluable tool for species delimita-
tion (Boury-Esnault et al., 2013; Gazave et al., 2013). At
present, the number of formally described sponge species is
comprised between 8000 and 9000, with an estimated 4000
species awaiting discovery and/or formal description
(Word Porifera Database, http://www.marinespecies.org/
porifera/).
In recent years, several contradicting hypotheses about
relationships at the basis of the metazoan tree and so
among nonbilaterian animals (Porifera, Cnidaria, Cteno-
phora and Placozoa) and among poriferan groups have
M
e Calcispongiae
t
a
z Demospongiae
o
a
(a)
Figure 1 Two main hypotheses currently proposed concerning nonbilaterian phylogenetic relationships: (a) paraphyly of sponges, (b) monophyly of
sponges. Modified from Ereskovsky et al. (2013). & Springer.
2 eLS & 2013, John Wiley & Sons, Ltd. www.els.net
been published (Dunn et al., 2008; Erwin et al., 2011;
Nosenko et al., 2013; Philippe et al., 2009, 2011; Pick et al.,
2010; Schierwater et al., 2009; Sperling et al., 2009; Wör-
heide et al., 2012). Although the majority of them support
the hypothesis of Porifera as a monophyletic group, sister
group of all other animals (commonly named Eumetazoa),
the debate is still open and the resolution of the base of the
metazoan tree remains among the most important open
questions in zoology. Sponges are now divided into 4 clades
(classes according to the Linnaean-ranked classification)
instead of the 3 recognised before 2009 (Gazave et al., 2012;
Figure 1). These 4 taxa – Hexactinellida, Demospongiae,
Calcarea and Homoscleromorpha – are distinguished by
different tissular, skeletal and developmental character-
istics, which will not be detailed here (main features are
summarised in Table 1).
The Calcarea are exclusively marine sponges and include
approximately 675 marine species (Word Porifera Data-
base). They have calcareous (calcite) spicular skeleton.
Spicules, little mineral pieces of variable size and shape, are
secreted in the extracellular space contrary to the intra-
cellularly formed siliceous spicules in other sponge taxa.
The skeleton is composed of free spicules, but some species
have a massive basal skeleton. Nevertheless, hypercalci-
fied, solid basal calcareous skeletons also occur in some
demosponges, but the crystalline structure is more often
aragonitic rather than calcitic as in Calcarea.
The Hexactinellida (or glass sponges) are also exclu-
sively marine sponges with siliceous skeleton. They are
largely restricted to the deep sea and cold waters. For this
reason, the diversity of this group (623 extant species
according to the World Porifera Database) is probably
Eumetazoa
M
e
Homoscleromorpha t
a
z
o
a
Hexactinellida (b)
 Porifera (Sponges): Recent Knowledge and New Perspectives

Table 1 Summary of the main characteristics of each clade of Porifera

Character Hexactinellida Demospongiae Calcarea Homoscleromorpha
Number of actual 601 6835 676 93
accepted species
Distribution All marine, mostly Marine, brackish and All marine All marine
deep waters freshwaters, all
depths
Nature of spicules Siliceous Siliceous when Calcium carbonate Siliceous when
present present
Types of spicules Megascleres/ Megascleres/ Megascleres/ No megascleres/
microscleres with microscleres with 1–4 microscleres with 1–4 microscleres
usually 6 or 3 axes axes axes distinction
Reproduction Viviparous Oviparous or Viviparous coelo- or Viviparous
viviparous more amphiblastula larvae cinctoblastula larvae
common
parenchymula larvae
Cell sheets The greater part of Cellular organisation Cellular organisation Cellular organisation,
soft tissue consists of flagellated
a single, pinacocytes, presence
multinucleate of a basement
syncytium membrane with type-
IV collagen
Types of aquiferous Syconoid, sylleibid or Leuconoid Asconoid, syconoid Sylleibid or leuconoid
system leuconoid pattern or leuconoid
Types of symmetry Radial Asymmetric or radial Asymmetric or radial Asymmetric
Source: According to Brusca and Brusca (2003); Word Porifera Database (http://www.marinespecies.org/porifera/); and Gazave et al., (2012).
# Elsevier.

underestimated. These sponges differ from the three other Ecology

classes in many aspects of their biology (Leys et al., 2007,
2011), mainly by a syncitial organisation instead of a
cellular one, and by 6-rayed spicules.
The Homoscleromorpha form the smallest group of
marine sponges (5100 described species). They can be
distinguished from others mainly by the presence of a
basement membrane (Boute et al., 1996; Ereskovsky et al.,
2009; Gazave et al., 2012) resulting in epithelial level orga-
nisation and tissue morphogenesis more similar to other
animals (Eumetazoa). The skeleton can be entirely absent.
When present, the spicules are peculiar tetractines (with 4
axes) and their derivatives secreted not only within scler-
ocytes as in demosponges but also within epithelial cells.
The Demosponges represent the most diversified taxa
not only in terms of species (85% of all sponge species)
but also regarding habitats conquered with noteworthy
adaptations to freshwater (temporary ecosystem in some
cases) and nutrient-depleted deep sea or caves (loss of
the aquiferous system and acquisition of macrophagy). The
mineral skeleton of siliceous spicules can be partially or
entirely replaced by an organic skeleton consisting of
abundant collagen fibres (spongin), as in bath sponges. The
skeleton can be totally absent, with a mesohyl containing
only abundant collagen fibrils. See also: Cambrian Explo-
sion; Cnidaria (Coelenterates); Ctenophora (Sea Walnuts
and Comb Jellies); Eukaryotes and Multicells: Origin;
Evolution of the Metazoan Extracellular Matrix; Placozoa
Sponges have a worldwide distribution and can be found
from tropics to polar regions. In some habitats, they can
represent the dominant macrobenthic organisms, con-
tributing up to 80% of the biomass. Their activities impact
on marine ecological processes at macro- and microlevels
by different ways:
1. Interactions with substrate: Boring sponges can exca-
vate calcareous substrates, such as mollusk shells, cor-
alline rhodophyta, coral reefs, etc. Some sponges are
thus actors of bioerosion (Carballo et al., 2012).
2. Trophic ecology: Most sponges feed on microorganisms
by a very effective filtration system. Exercising little
selection (except size limit imposed by ostia diameter),
they pump a large volume of water, cycling a volume
equivalent to their body volume every 7 s (Bergquist,
1978; Reiswig, 1971). Their presence thus impacts the
microbial abundance and diversity. Moreover, many
species of unicellular eukaryotes, archaea and eubacteria
are hosted symbiotically by sponges (for illustration:
pinkish colour visible on Figure 3e corresponds to the
presence of cyanobacteria), which can be considered as a
reservoir of microbial diversity. Some of them hosting
symbiotic cyanobacteria can become net primary pro-
ducers and contribute to oxygen production (Webster
and Taylor, 2012). A few sponge species that live in deep

eLS & 2013, John Wiley & Sons, Ltd. www.els.net 3

 Porifera (Sponges): Recent Knowledge and New Perspectives
sea, a food-poor environment, have become carnivores
and prey mainly on small crustaceans (Vacelet and
Boury-Esnault, 1995; Vacelet and Duport, 2004).
3. Interactions with other organisms: Sponges also often
host small metazoans or their larvae (arthropods,
nematodes, annelids, echinoderms and cnidaria) for
which they offer shelters.
The defences of sponges against pathogens, competitors
and predators (such as echinoderms, mollusks, fishes, etc.)
mainly consist in the presence of spicules (mechanic
defence) and production of toxic secondary metabo-
lites (chemical defence). Sponge secondary metabolites
obviously present a defensive role against predation,
infections and fouling. Indeed, they exhibit antibacterial,
antifungal, antiplasmodial, antihelminthic, antiamoebitic,
antileishmanial as well as anti-inflammatory and anticancer
activities. Among the chemical compounds isolated so far
from sponges (more than 5000), 800 present antibiotic
properties, thus opening up future medical applications
(Darah et al., 2011; Torres et al., 2002; Touati et al., 2007).
Moreover, these chemicals play an important role in
Exhalant canal
Exhalant pore
(Osculum)
Endopinacoderm
Exopinacoderm
Inhalant canal
Inhalant pore
(Ostium)
Mesohyl
(a)
Osculum
Osculum
Ascon Sycon
(b)
Figure 2 (a) General organisation of sponges. (b) The three main levels of complexity of the aquiferous system. Modified with permission from Philippe
et al. (2009). & Elsevier.
4 eLS & 2013, John Wiley & Sons, Ltd. www.els.net
competition for space, indeed some of them are lethal
weapons to fight against other sessile and encrusting inver-
tebrates (such as other sponge species, cnidarians, bryozo-
ans, ascidians, etc.).
General Organisation of the Body and
the Aquiferous System
The cell layer responsible for the main delimitation of the
body is called pinacoderm, which is composed of flattened
cells named pinacocytes (baso-, exo- and endopinacocytes
according to their position). Internally, the pinacoderm
lines (inhalant) canals that carry the water to the filtering
cells – the choanocytes (often organised in chambers,
see Figure 2). Choanocytes (forming a cell layer named
choanoderm) are highly specialised collar cells. Water flow
is ensured by the beating of choanocyte flagella while the
microvilli of the collar trap unicellular organisms that will
be phagocytised. The filtered water leaves the sponge
by (exhalant) canals lined by pinacoderm. Choanocytes
Flagellum
Choanocyte
Microvillosities
Spicule
Choanocyte chamber
Osculum
Choanoderm
Leucon

account for 80% of organic carbon uptake; nevertheless,
surface- and canal-lining pinacocytes can phagocytise
particles directly. The canals and choanocyte chambers
network form together the aquiferous system. The aqui-
ferous system can be more or less complex, leading to a
peculiar terminology. Figure 2 provides a schema of the
general organisation of a sponge (Figure 2a) and the main
types of aquiferous systems (asconoid, syconoid and
leuconoid used for species identification and classification,
Figure 2b). The water flow can be regulated, in response
to environmental changes, by dilatation or restriction
of inhalant pores (ostia), canals and exhalant pores
(oscula) diameter. In addition, active pulsatile contrac-
tions observed by time-laps imaging are supposed to par-
ticipate to the efficiency of pumping (Nickel et al., 2011).
The mechanisms of signal transduction mediating per-
ception and regulation of pumping are nevertheless still
misunderstood, even if neuromediators are supposed to be
involved (for review see Renard et al., 2009). One exception
is worth noting: the aquiferous system and its choanocytes,
long considered as characteristic of sponges, are absent
(maybe lost) in most carnivorous sponges.
Between pinacoderm and choanoderm, a fibril-rich,
looser and less organised layer is found: the mesohyl, which
contains various types of cells (the number of which depends
on the group considered), notably totipotent cells called
archaeocytes (see Cell Plasticity, Stem Cells and Cell
Renewal), cells involved in mineral and/or organic skeleton
synthesis (such as sclerocytes, spongocytes and lophocytes),
cells supposed to be implicated in chemical synthesis
and exocytosis (granular and spherulous cells) as well as
gametes, skeleton elements and symbiotic microorganisms.
Mineral skeleton, when present, is either siliceous or
calcareous, usually in the form of spicules that have a
bewildering range of shapes, sizes and patterns of organi-
sation (at the basis of species identification and classifica-
tion). The larger elements (megascleres) maintain the
overall form of the sponge, whereas tiny spicules (micro-
scleres) line the surfaces or are dispersed without order.
This part can be concluded saying that as sponges are
devoid of organs, their individual cells or cell layers ensure
vital functions. Feeding involves choanoderm and pina-
coderm; most of other activities involve mesohyl cells, for
example, the synthesis of organic and mineral skeleton.
See also: Evolution of the Metazoan Extracellular Matrix
Cell Plasticity, Stem Cells and Cell
Renewal
Regarding the low number of cell types in sponges (ranging
from 4 to 12 (Ereskovsky, 2010)), some authors consider
that one cell type may have several functions even if con-
crete proof sometimes lacks: for example, choanocytes are
obviously involved in nutrition and reproduction (see
‘Reproduction and Development’) but may also play sen-
sorial roles (Jacobs et al., 2007; Gazave et al., 2008; Renard
eLS & 2013, John Wiley & Sons, Ltd. www.els.net
Porifera (Sponges): Recent Knowledge and New Perspectives
et al., 2009) and pinacoderm is involved in body lining and
nutrition but also ensures contractility (Nickel et al., 2011).
In addition to cell plurifunctionality, the functional plas-
ticity of sponges also relies on a high capacity of cell
transdifferentiation.
Indeed, as in other metazoans, cell renewal involves
pluripotent cells. In text books, archaeocytes are generally
considered as demosponge stem cells based on morpholo-
gical observations, and this was recently supported by
molecular data (Funayama, 2013). But in addition, it was
shown that choanocytes also express stem cell gene mar-
kers, which suggests that these specialised cells retain
pluripotency. This is consistent with the fact that they are
often involved in gamete formation by transdifferentiation
(Funayama, 2013).
This cellular plasticity may explain the sponge’s great
capacity to regenerate or reaggregate, which offers very
interesting technical possibilities to study cell differentia-
tion processes. Sponges possess remarkable reconstitutive
and regenerative abilities ranging from not only the com-
mon wounds cicatrisation to body part regeneration but
also to functional body regeneration from dissociated
cells. Indeed, sponge cells can be dissociated into cell sus-
pensions (chemically or mechanically) and then, under
appropriate ionic conditions, cells can reaggregate and
reconstitute either organised structures named primorphs
or, in some cases, entirely functional individuals. If mix-
tures of cells of different species are realised, they recon-
stitute separately, proving that cell recognition complexes
also exist in these organisms (also shown by graft experi-
ments; for review see Simpson, 1984). See also: Adhesive
Specificity and the Evolution of Multicellularity
Reproduction and Development
Reproduction and development was more largely studied
in Demospongiae and Calcarea, but nevertheless they
show a great diversity of strategies. Indeed, both sexual and
asexual reproductions are commonly used by sponges,
with a large diversity of reproductive processes.
Like in other metazoans, sexual reproduction involves
gametogenesis processes leading to the formation of ani-
sogamic gametes with typical mobile spermatozoids and
oocytes containing vitellus. Sponges can be either gono-
choristic or hermaphroditic (in this case both protandry
and protogyny have been described), hermaphroditism
being considered as more frequent. Gametes are not
produced in gonads but result either from archaeocytes
differentiation or choanocytes transdifferentiation. Ferti-
lisation can occur either internally in the mesohyl or
externally in the water column. In the first case, sperma-
tozoids enter the oocyte coating by the aquiferous system.
An impressive mechanism has been described in Calcarea:
spermatozoids are captured by choanocytes that dediffer-
entiate to become carrier cells transporting them to
oocytes. In all cases, the obtained zygote enters in devel-
opmental processes leading to a mobile, ciliated larval
5
 Porifera (Sponges): Recent Knowledge and New Perspectives

stage characteristic of each sponge groups. After a short
free life (2 h to 4 days), the larva settles on a selected sub-
strate and metamorphoses to give rise to a sessile filtrating
adult (Ereskovsky, 2010). For some species, the embryos of
different stages develop in brood chambers until they reach
a certain size and then disperse.
Asexual reproduction can occur by external (budding)
or internal (gemmules) tissular reorganisation processes as
well as fragmentation. Formation of buds, which subse-
quently detach, is a common asexual process in marine
sponges, whereas the formation of gemmules, a state of
quiescence consisting in archaeocyte aggregates encapsu-
lated within a spongin coat, characterises most of
freshwater sponges. For the latter case, on hatching, all
adult cells derive from the archaeocytes, an excellent
illustration of their totipotency. Gemmules formation
sometimes also occurs in some shallow-water marine
Demospongiae
sponges (Simpson, 1984). They show impressive resistance
features allowing the colonisation of temporary waters and
high possibilities of adaptation to fluctuating environments
(Schill et al., 2006).
Thereby, according to the few well-described sponge
life cycles, it seems that sponges often use both asexual
and sexual reproduction during their lifespan. See also:
Reproduction and Life Cycles in Invertebrates
New Insights and Perspectives on
Early Animal Evolution
Because of their morphoanatomical simplicity, Porifera
have not been considered as true animals for a long time,
sometimes regarded as colonial organisms and a step

(a) (b) (c)

Calcarea

(d) (e) (f)

Homoscleromorpha
Hexactinellida

(g) (h) (i)

Figure 3 Pictures illustrating the diversity of forms, colours and sizes in Porifera, with a few chosen examples from the four clades. (a) Aplysina cavernicola,
(b) Ephydatia fluviatilis, (c) Xestospongia muta, (d) Sycon ciliatum, (e) Clathrina contorta, (f) Leucosolenia complicata, (g) Rosella nuda, (h) Oopsacas minuta and
(i) Oscarella species (Oscarella tuberculata and Oscarella lobularis). & Elsevier.

6 eLS & 2013, John Wiley & Sons, Ltd. www.els.net

 Porifera (Sponges): Recent Knowledge and New Perspectives

Hexactinellida Demospongiae Calcarea Homoscleromorpha Eumetazoa

3 3
4 4

3
4

2 3
4 5

1
3

Figure 4 Possible scenarios of character evolution during animal history, according to the (currently most accepted) hypothesis of monophyly of sponges.
Lines represent acquisition of characters, crosses represent losses of characters, red or green colours represent alternative scenarios for a same character.
Characters are coded by the following numbers: 1: multicellularity; 2: aquiferous system; 3: basement membrane and true epithelia; 4: siliceous spicules;
5: neurones, muscle cells and digestive system. & Elsevier.

between uni- and multicellular organisms. Molecular,
phylogenetic and embryological data accumulated since
the end of the twentieth century have closed this debate
showing that they are bona fide Metazoa. The more
molecular (transcriptomic and genomic) data are acquired,
the more the complexity of sponge genome content raises
exciting questions. Indeed, most of key transcription factor
families and main signalling pathways required for eume-
tazoan development and body patterning are present
in sponges, suggesting that the last common ancestor of
Metazoa (named Urmetazoa) already owned a complex
molecular toolkit (Ereskovsky et al., 2013). The next step in
progress to be reached is determining the roles of all these
genes in sponges and comparing them to eumetazoan in
order to propose evolutionary scenarios of gene function
evolution and try to correlate them with body plan evolu-
tion (Figure 3 proposes scenarios of character acquisitions).
So far, only two whole sponge genomes are at present
publicly available and few others are in progress. The
recent rising interest for Porifera is expected to allow a
better understanding of early animal evolution by the
growing amount of cellular, molecular and functional
data: Was Urmetazoa a sponge-like simple organism or, in
contrast, a more complex animal; the simplicity of sponges
then being the result of a secondary simplification, as
observed several times in animal evolution (e.g. cirripeda
and tunicates)? (Figure 4; Nielsen, 2008; Philippe et al.,
2009; Wörheide et al., 2012). See also: Animals and Their
Unicellular Ancestors; Nuclear Receptor Genes: Evolu-
tion; Origin of Bilaterian Hox Patterning System
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Further Reading
Funayama N (2010) The stem cell system in demosponges:
Insights into the origin of somatic stem cells. Development,
Growth & Differentiation 52(1): 1–14.
Leys SP (2004) Gastrulation in Sponges. In: Stern CD (ed.) Gas-
trulation: From cell to embryo, pp 23–32. New York: Cold
Spring Harbor Lab Press.
Manuel M (2009) Early evolution of symmetry and polarity in
metazoan body plans. Comptes Rendus Biologies 332(2–3):
184–209.