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Integrated Principles of Zoology

Eighteenth Edition

Chapter 12

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 Sponges and Placozoans

© McGraw Hill ©William C. Ober/Medical Scientific Illustration 2

 The Origins of Multicellularity

Cells are the elementary units of life.

• Nature’s experiments with larger organisms without

cellular differentiation are limited.
• Large, single celled marine algae are rare.

Increasing the size of a cell causes problems of exchanging

molecules with the environment.

Multicellularity prevents surface-to-mass problems as smaller

units greatly increase surface area for metabolic activities.

• Highly adaptive toward larger body size.

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 Multicellular Sponges

Sponges are the simplest multicellular animals but their cell

assemblages are distinct from other metazoans.
• Sponges have cells embedded in an extracellular matrix
supported by a skeleton with needle-like spicules and
protein.
Sponges neither look like nor behave as animals but
molecular evidence demonstrates that they are
phylogenetically grouped with animals.

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 Origin of Animals (Metazoa)

Eukaryotic cells evolved and diversified into many lineages

that led to modern day descendants.

• Includes all unicellular eukaryotes, colonial and

multicellular plants, animals, and fungi.

• Multicellular organisms were collectively called metazoans

and are now also termed “animals.”

• Metazoans are placed in the Opisthokont clade which

include fungi, choanoflagellates, and other groups.

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 Choanoflagellates
Choanoflagellates are solitary or colonial aquatic
eukaryotes.
• Each cell has a flagellum surrounded by a collar of
microvilli.
• Flagellum beats and draws water into collar where
the microvilli collect particles like bacteria.
• Most are sessile// but one species attaches to
floating diatom colonies and feed midwater.
• Strongly resemble sponge feeding cells called
choanocytes.

• Choanoflagellates and choanocytes may share a

common ancestor.

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 Choanoflagellates and Metazoans

Evidence of common ancestry.

• Proteins used by colonial choanoflagellates for cell
communication and adhesion are homologous to those
that metazoans use in cell-to-cell signaling.
Evidence against common ancestry.
• Choanocytes are not part of the developmental pathway of
sponges; they appear only in adult sponges.
• Collar cells appear in a few other animal taxa; if this trait is
ancestral then it has been lost in most taxa.

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 Approaches to Animal Origins

Compare sponge genomes to those of other animals.

• Shared features would have been inherited from ancestor
of animals.
Sponges today have simple bodies, and therefore would be
expected to have simple genetic structure.
• Sponge genome contains elements that code for
regulatory pathways of more complex metazoans.
• Have proteins that code for spatial patterning that specify
anterior and posterior pole of larvae.
• Therefore, modern sponges may be less morphologically
complex than their ancestors.
Phylum Placozoa also exhibits puzzling features.
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 Phylum Porifera: Sponges

General features of sponges.

• Mostly sessile.
• Body designed for efficient aquatic filter feeding.
• Porifera means “pore-bearing”; sac-like bodies are
perforated by many pores.
• Use flagellated “collar cells”, or choanocytes, to move
water to bring food and oxygen while removing wastes.
• Most of the 8600 sponges are marine, found in all seas
and all depths, while few live in brackish water and 150
live in fresh water.

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 Growth Habits and Forms of Sponges

Figure 12.1 Some growth habits and forms of sponges.

Access the text alternative for slide images.

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 Sponge Collar Cells

Figure 12.2 Sponge choanocytes have a collar of microvilli surrounding a

flagellum.
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 More Sponge Features

Sponges vary in size from a few millimeters to over 2 meters in

diameter.
Many species are brightly colored because of pigments in dermal
cells.
Embryos are free-swimming while adult sponges always attached.
Growth form varies.
• Some appear radially symmetrical but many are irregular in
shape.
• Some stand erect, some are branched, and others are
encrusting.
• Growth patterns depend on shape of substratum, direction of
water, speed of flow, and availability of space.
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 Interactions

Many animals such as crabs, nudibranchs, fish, and other

species live as commensals or parasites in or on sponges.
Sponges can also grow on a variety of other living organisms
with some crabs using sponges for camouflage and
protection.
Sponges and microorganisms living on them often have a
noxious odor and produce a variety of bioactive compounds.
• Certain sponge extracts have manifested medical and
pharmaceutical effectiveness.

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 Skeletal Structure

Skeletal structure of a sponge can be fibrous and/or rigid

consisting of calcareous or siliceous spicules.
Fibrous portion comes from collagen protein fibrils in
intercellular matrix.
• There are several types of collagen, which vary in
chemical composition; sponges contain spongin.
Composition and shape of spicules are the basis of
classification.
• Modern materials science view spicules for possible
nanoparticle products.
Simplistic exterior of sponges often mask their chemical and
functional sophistication.
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 Spicule Forms

Figure 12.3 Diverse forms of spicules that support a sponge body.

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 Poriferan Classification

Sponges date to early Cambrian, maybe even Precambrian.

Traditionally grouped in three classes based on spicules and
chemical composition.
• Calcispongiae: calcium carbonate spicules with one, three,
or four rays.
• Hexactinellida: glass sponges with six-rayed siliceous
spicules.
• Demospongiae: siliceous spicules around an axial
filament, spongin fibers, or both.
Class Homoscleromorpha identified as sponges without a
skeleton, or with siliceous spicules without an axial filament.

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 Cladogram of Poriferans

Figure 12.4 Cladogram depicting evolutionary relationships among the

four classes of sponges.
Access the text alternative for slide images.

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 Feeding in Sponges

Sponges feed by collecting suspended particles from the

water pumped through internal canal systems.
• Many small incurrent pores (dermal ostia) in pinacoderm
(outer layer of cells).
• Water is directed past the choanocytes, which are
flagellated collar cells that keep the current flowing via
beating of flagella.
• Microvilli in the collar trap and phagocytize food particles
that pass by.
Efficiency of food capture is dependent on water movement
through the sponge body.

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 Food Particle Consumption

Sponges non-selectively consume food particles (detritus,

plankton, and bacteria).
• The smallest particles (80%) are taken into choanocytes
by phagocytosis.
• Protein molecules may be taken in by pinocytosis.
• Two other cell types, pinacocytes and archaeocytes,
facilitate feeding.
• Dissolved nutrients can also be absorbed by sponges.

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 Sponge Body Structures

Figure 12.5 Three types

of sponge structure.

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 Asconoid Sponge Body

Simplest body organization.

• Small and tube-shaped to allow water to flow directly
across cells so no “dead space.”
• Choanocytes are in a large internal chamber, the
spongocoel.
• Choanocyte flagella pull water through the pores and extract food
particles.
• Used water is expelled through a large single osculum.

• All asconoids are in class Calcispongiae.

• For example, Leucosolenia sp. and Clathrina sp.

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 Asconoid Sponge

Figure 12.6 Clathrina canariensis (class Calcarea) is a common

Asconoid on Caribbean reefs.

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 Syconoid Sponge Body

Syconoids resemble asconoids, but larger with a thicker,

more complex body wall.
• Body wall is folded outwards with choanocyte-lined radial
canals that empty into spongocoel.
• Water enters through dermal ostia and move through tiny
openings (prosopyles) into the radial canals.
• Food is ingested by choanocytes and used water is
pumped through internal pores called apopyles then
outwards via osculum.
• Spongocoel is lined with epithelial cells rather than
choanocytes as in asconoids.

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 Syconoid Evolution and Distribution

Developmental evidence of being derived from asconoid

ancestors.
• Syconoids pass through an asconoid stage in
development but do not form highly branched colonies.
• Flagellated canals form by evagination of the body wall.
Syconoid body plan is not homologous among all sponges
that have it.
• Classes Calcispongiae and Hexactinellida have syconoid
species (ex: Sycon sp.)

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 Syconoid Body Wall

Figure 12.7 Cross section through wall of sponge Sycon sp., showing
choanocytes in canals within the wall but do not line spongocoel.

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 Leuconoid Sponge Body

Most complex and largest, with more food-collecting regions.

• These regions have choanocytes lining in small chambers
that effectively filter all water present.
• Clusters of flagellated chambers are filled from incurrent
canals and discharge to excurrent canals which lead to
osculum.
• After food is removed, used water is pooled to form an exit
stream that leaves through an exit pore at very high
velocity.
• This high rate of exit flow prevents the sponge from re-filtering used
water and wastes.
• Most sponges are leuconoid type.
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 Leuconoid Evolution and Distribution

The leuconoid system has high adaptive value to efficiently

meet high food demands of larger body size.
• Highest proportion of flagellated surface per volume of cell
tissue.
• More collar cells can filter more particles.
• Water flow slows inside due to greater surface area in the
chambers.
• Large sponges filter 1500 liters of water per day for
maximum food collection.
The leuconoid system has evolved independently many
times in sponges.

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 Leuconoid Sponge

Figure 12.8 This orange demosponge, Mycale laevis, often grows

beneath plate-like colonies of the stony coral Montastrea annularis.
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 Types of Cells in the Sponge Body

Sponge cells are arranged in a gelatinous extracellular

matrix called mesohyl or mesenchyme.

• The connective “tissue” of sponges found in fibrils, skeletal

elements, and amoeboid cells.

Absence of organs requires that all fundamental processes

occur at the individual cell level.

• Respiration and excretion via diffusion and water

regulation via contractile vacuoles in the archaeocytes and
choanocytes.

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 Sponge Activity and Response

Visible activities seen in sponges include.

• Slight alterations in shape, local contraction, and
propagating contractions.
• Closing and opening of incurrent and excurrent pores.
Sponges can close their osculum due to heavy sediment
load.
• Movements occur very slowly but they suggest a whole
body response in organisms lacking complex organization
above the cellular level.
• Apparently excitation spreads from cell to cell by
mechanical stimuli and signaling molecules such as
hormones or via electrical impulses.
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 Types of Sponge Cells

Figure 12.9 Small section through sponge wall, showing four types
of sponge cells.

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 Choanocytes

Choanocytes are ovoid cells.

• One end embedded in mesohyl; exposed end has
flagellum surrounded by a collar.
• Collar consists of microvilli connected to each other by fine
microfibrils.
• Forms a fine filtering device to strain food.
• Particles too large to enter collar are trapped in mucous
and slide down to base to be phagocytized.
• Food is passed to archaeocytes for intracellular digestion
with no need for gut cavity.

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 Sponge Cells Trapping Food

Figure 12.10 Food trapping by sponge cells. (A) Cutaway section of canals
showing direction of water flow. (B) Two choanocytes, and (C) Structure of the
collar.

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 Archaeocytes

Archaeocytes are amoeboid cells with many functions that

move about in the mesohyl.
• Phagocytize particles in the pinacoderm.
• Receive particles for digestion from choanocytes.
• Can differentiate into many other more specialized cell
types.
• Sclerocytes: secrete spicules.

• Spongocytes: secrete sponging.

• Collencytes: secrete fibrillar collagen.
• Lophocytes: secrete large amounts of collagen.

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 Pinacocytes

Pinacocytes are thin, flat, and epithelial-like cells.

• Cover the exterior and interior surfaces of sponges almost

like real tissues.

• Form pinacoderm with a variety of intercellular junctions

but no basal membrane in most sponges.

• Ingest food by phagocytosis and are contractile to regulate

surface area of sponge.

• Form myocytes, circular bands around oscula, that help

regulate flow of water.

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 Cell Independence: Regeneration and Somatic
Embryogenesis

Sponges have a great ability to regenerate lost parts and

repair injuries.
Complete reorganization of the structure and function of
participating cells or bits of tissue occurs in somatic
embryogenesis.
Process of reorganization differs in sponges of differing
complexity.
Regeneration following fragmentation is one means of
asexual reproduction.

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 Asexual Reproduction

Fragmentation.
• Sponge breaks into parts that are capable of forming a
completely new sponge.
Bud formation.
• External buds.
• Small individuals that break off from parents that have reached a
certain size.

• Internal buds or gemmules.

• Formed by archaeocytes that collect in mesohyl.
• Coated with tough spongin and spicules that can survive harsh
environmental conditions.

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 Gemmulation

When parent sponge dies, gemmules survive and remain

dormant during the harsh situations.
Live cells within gemmules escape through special opening
called micropyles and develop into new sponges.
Gemmulation is a adaptation to changing seasons and for
colonization of new habitats.
Gemmules are controlled by weather, internal chemicals, and
by remaining inside the parent sponge.

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 Gemmule Cross Section

Figure 12.11 Section through a gemmule of a freshwater sponge

(Spongillidae).
Access the text alternative for slide images.

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 Sexual Reproduction

Most sponges are monoecious (both male and female sex

cells in one body).
Gametes develop from choanocytes or archaeocytes.
Most sponges are viviparous.
• Zygote is retained within parent and provided with
nourishment until it is released as a ciliated larva.
• One sponge releases sperm which enter the pores of
another sponge.
• Choanocytes that phagocytize the sperm and transport it
through mesohyl to oocytes to form zygotes.

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 Poriferan Larvae

Some sponges are oviparous.

• Oviparous sponges release both sperm and oocytes into
water for external fertilization.
The free-swimming larva of most sponges is a solid-bodied
parenchymula larvae.
• Six other larval forms also exist.
• Outwardly directed flagellated cells of the parenchymula
become choanocytes.

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 Unique Developmental Pattern

Found in Calcispongiae and some Demospongiae.

• Hollow stomoblastula develops with flagellated cells oriented
toward the interior.
• Blastula then turns inside out (inversion) and the flagellated
cells now turn outside.
• Small flagellated cells or micromeres located at anterior end
while larger non-flagellated macromeres located at posterior
end.
• Macromeres overgrow invaginating micromeres during
metamorphosis and settlement.
• Micromeres become choanocytes, archaeocytes, and
collencytes while macromeres give rise to pinacoderm and
sclerocytes.
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 Typical and Unique Development Patterns

Figure 12.12 (A) Development of demosponges, and (B) Development of

the calcareous syconoid sponge Sycon sp.

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 Class Calcispongiae

Calcareous sponges with spicules of calcium carbonate.

Spicules are straight (monaxons) or have three or four rays.
Most are small with tubular or vase shapes.
Many are drab in color, but some are bright yellow, green,
red, or lavender.
Leucosolenia and Sycon are commonly studied marine
shallow-water genera.
Asconoid, syconoid, and leuconoid body forms.

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 Class Hexactinellida

Also known as Class Hyalospongiae.

Glass sponges with six-rayed spicules of silica bound
together to form network.
Nearly all are deep-sea forms.
Most are radially symmetrical with vase or funnel shaped
bodies attached by stalks of root spicules onto the substrate.
Have syncytial cell structure.
• Many nuclei within a large cell.
• Produced by the fusion of many cells or division of nuclei
without dividing the cytoplasm.

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 A Glass Sponge

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 Class Demospongiae

Contains 95% of living sponge species and most large

sponges.
Spicules are siliceous but not six rayed and may be absent
or bound together by spongin.
Leuconoid body form for all species.
All marine except for Spongillidae, the freshwater sponges.
Marine demosponges are highly varied in color and shape,
with some growing to several meters in diameter.

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 Freshwater Demosponges

Widely distributed in well-oxygenated ponds and streams.

They encrust plant stems and submerged wood.
Look like wrinkled scum, pitted and porous with brown and
green colors.
Flourish in summer and in early autumn.
Reproduce sexually, but existing genotypes may also
reappear annually from gemmules.
Sponges die by late autumn and asexually release
gemmules to prepare for next year’s population.

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 Marine Demosponges

Figure 12.15 Marine Demospongiae on Caribbean coral reefs. (A)

Pseudoceratina crassa, (B) Aplysina fistularis, and (C) Monanchora unguifera

© McGraw Hill (a,c): ©Larry Roberts/McGraw-Hill Education b:©William C. Ober/Medical Scientific Illustration 50
 Class Homoscleromorpha

Mostly marine with a variety of colors but live in cryptic

habitats.
Generally found near shore but have deep water forms.
Separated from Demospongiae due to presence of true
basement membrane under pinacoderm or extracellular
matrix.
Also have adherens cell junctions that from true tissues
unlike other sponges.
Divided into two clades based on absence or presence of
spicules.

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 Phylogeny

Sponges appeared before the Cambrian.

• Two groups of calcareous sponge-like organisms were in early
Paleozoic reefs.
Sponges share many traits with other animals and are considered
sister taxon.
• Proteins for cell adhesion and cell-signaling are homologous to
other animals.
• Some sponges have basement membrane with collagen and
adherens junctions with cadherin molecules that connect
epithelial cells.
• Sponge have blastula and some form gastrula stages like many
animals.
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 Adaptive Diversification

Poriferans are a highly successful group with thousands of

species in diverse habitats.
Diversification centers on their unique water-current system
and its degree of complexity.
New feeding mode has evolved for sponges found in deep
water caves with low nutrients.
• Illustrates the non-directional nature of evolution.

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 Unique Deep Water Sponges

Many tiny hook-like spicules cover highly branched body.

Spicule layer can entangle the legs of crustaceans that come
near sponge.
Filaments of the sponge body grow over prey, slowly
enveloping it and later digesting it.
Most of the group are carnivores and not suspension
feeders.
Some have symbiotic methanotrophic bacteria.
Contain siliceous spicules, but lack choanocytes and internal
canals making them very different than regular sponges.

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 Carnivorous Harp Sponge

Figure 12.16 The carnivorous sponge, Chondrocladia lyra, is commonly

called a “harp sponge.”

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 Phylum Placozoa

Proposed by K. G. Grell (1971) based on a single species,

Trichoplax adhaerens.
• Tiny (2 to 3 mm) marine form that is plate-like and has no
symmetry.
• No major organs, no muscular or nervous system.
• Lacks basal lamina beneath epidermis and no extracellular
matrix but has genes for it.
• Body has dorsal epithelium to cover cells and thick ventral
epithelium of monociliated cells and nonciliated gland cells.
• Space between the epithelia contain multinucleated fibrous
“cells” within a contractile syncytium.

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 Placozoan Behavior

Placozoans glide over food, secrete digestive enzymes, and

absorb nutrients.
Divide asexually and produce “swarmer” stages by budding.
• No sexual stages have been seen but have isolated eggs
in the laboratory.
Considered diploblastic.
• Dorsal epithelium represents ectoderm.
• Ventral epithelium represents endoderm.
• Genetic studies support these homologies.

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 Placozoan Structure

Figure 12.17 (A) Trichoplax adhaerens is a marine placozoan, and (B)

Section through Trichoplax adhaerens, showing histological structure.

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