Ch- 4: INTRO. TO METAZOA {R.D.

Barnes}

METAZOA
● Evolution from protozoa
● Protozoans are unicellular eukaryotic organisms that often form multicellular
colonies → evolutionary door open
● 1 million- 30 million sp. In 29 major taxa (phyla)

GROUND PLAN:
Metazoans are
● multicellular,
● heterotrophic,
● motile eukaryotes,
● body polarised along ant-post. locomotory axis,
● generally large than protozoans

CELLS, TISSUES, & SKELETONS

● Metazoans body- functionally specialised cells
● Protozoans- all func. in a single cell
● Metazoans bound together in specialised layers= tissues
● (minimally 2 tissues, at least 1 more than any colonial protozoans. Eg-
protozoan colony Volvox - const. The sole tissue layer of the body; 2
tissue eg- Choanoflagellata - sister taxon of Metazoa)
● 2 types of tissues- Epithelial & Connective- maybe primary(original)
metazoan tissues
● Epithelium-
○ cells that adjoin one another to form a sheet that covers the body or
lines an internal cavity;
○ on its ext. surf. a secreted extracellular matrix (ECM),or cuticle
○ Cuticle- consists of little more than a surface coat of glycoprot.
○ Epi. cells- secrete and rest on a basal lamina, a thin, dense, fibrous
ECM
○ Bounded by cell-adhesion molecules
○ Classes- Integrins bind cells to the basal lamina; Cadherins to each
other
○ Commonly ciliated
○ Monocilated- primitive eg- in some apomorphic(specialised trait
unique to a gp. Or spe.) groups and in Choanoflagellates
○ The Epithelium is an unbroken layer of cells that provides a
physiological barrier between the interior of the organism and the
 exterior environment. It is anchored between a basal lamina below
and a secreted, protective cuticle above. Secretory cells are
interspersed among the monociliated cells. The connective-tissue
layer (or compartment) consists of a proteoglycan gel containing
collagen fibers and individual connective-tissue cells.
○ Both the cuticle and internal ECM function as a skeleton

Hypothetical Metazoan with 2 primary metazoan tissues: monociliated, ciliated

epithelium overlying connective tissue
● Connective tissue-
○ widely separated, non-adjusting cells in ECM
○ ECM- less voluminous, const. proteoglycan gel with embedded prot.
fibres; most common prot. - collagen (in humans, amount for
1-quarter of all prot. in body)
○ collagen fibres + connective tissue have skeleton function
○ ECM- provide attachment sites for cells, cell moving pathways,
environ. for cell differentiation → imp. role in organising &
maintaining tissues of metazoan body
● Skeleton- str. that supports the body & transmits force of muscular
contraction
○ provide protection from- predation, injury, infection, or environ.
challenge
○ metazoan- tissue level str.
○ metazoans- have exo, endoskeleton or both
○ Exoskeleton- thickened cuticle that becomes rigid by chemically
cross-linking its
■ proteins- as in insects
■ adding minerals to org. matrix- as in crabs
■ chemical cross-linking- as in cartilage
■ mineral secretion- as in bone; to prod. endoskeleton
REPRODUCTION & DEVELOPMENT
● both sexual & asexual repro.
● Asexual/ Colonal repro.
○ Fragmentation- body breaks up irregularly into several pieces
○ Fission- more orderly division along longitudinal or transverse axis
○ following both- regenerate missing parts
○ Budding- differentiation of a daughter individual before it detaches
from parent body
○ Parthenogenesis (=virgin birth)- dev. of an individual from an
unfertilized egg or other totipotent cell
● Sexual repro.
○ are diploid org. in which meiosis is restricted to the form. of haploid
gametes; sperm & egg; both specialised, polar cells
○ Reunion= Fertilization - restores diploidy and produces a polarized
zygote, or fert. egg
○ polar axis of zygote runs b/w animal & vegetal poles
○ in most cases, the animal pole corresponds to the egg’s apical end,
the former site of a flagellum & microvillar collar that regressed
(revert) during differentiation of the egg
○ zygote- divides by mitosis = prod. a multicellular embryo
○ early divisions of zygote= cleavages & resulting cells =
blastomeres
○ 1st 2 div. planes are || to animal-vegetal axis(meridional) and entire
zyg. cleaved into 2 & then 4 blastomeres = such Complete cleavage
is Holoblastic cleavage
● Eggs- various amts. of yolk distributed
○ yolk inc. the size of egg and tends to interfere with cleavage, both
the amt. & distribution of yolk affect cleavage pattern & embryo
shape
○ Microlecithal eggs- small cells, little yolk distributed evenly
throughout the cytoplasm; undergoes holoblastic cleavage; prod.
blastomeres of equal size (equal cleavage)
○ Mesolecithal eggs- medium size, moderate amt. of yolk restricted
to vegetal hemisphere; undergoes holoblastic; unequal cleavage
■ blastomeres are smaller= micromeres
■ macromeres
○ Macrolecithal eggs- large, v. yolky, low area-to-vol. ratios, no
holoblastic cleavage, incomplete cleavage= meroblastic occurs
 ■ initially cleavage furrow memb. incomplete, do not fully isolate
cytoplasm from yolk mass
■ later complete memb. separate thin cap of cells= blastodisc
from yolk; provisioned by diffusion of yolk nutrients & gases
○ Macrolecithal eggs & meroblastic cleavage occurs in squids,
octopuses, certain fishes, birds, & reptiles.
○ Superficial cleavage- incomplete cleavage- macrolecithal eggs of
insects & other arthropods.
○ early embryos form Blastula- 1 cell thick cell ball, either
hollow(coeloblastula); or solid (stereoblastula)
○ after cell div. → blastomere move into interior of blastula =
gastrulation [early dev. process in which the embryo transforms
from a 1-D layer of epithelial cells(blastula) & reorganises into
multilayered & multidimensional str. called gastrula]; from 2 layers=
gastrula
○ Gastrulation-1st event in morphogenesis (conversion of less
uniform embryo to complex, multilayered adult)
○ in most ani.- primary germ layers (ecto & endoderm) formed by
gastrulation
○ b/w ecto & endoderm is gelatinous layer of ECM= blastocoel
○ embryonic precursors of adult tissues- ecto, endoderm & blastocoel
○ dev.-
■ ectoderm becomes outer (epidermis)
■ endoderm- inner (gut) layers
■ blastocoel is dev. precursor of connective tissue
○ after the gastrula stage, dev. embryo inc. in complexity (forms
organs) and adopts typical anatomy of spe.
● events unfold during dev. period from zygote to adult constitutes its
ontogeny(=origin of spe.)
○ has time course
○ adaptations in dev. affect reproductive success of adults, natural
selection modified ontogenetic & adult stages
○ eg- larva (a ontogenetic modification)
● larva- dev. stage of distinctive form
○ persists for a period of time
○ lives independently of adult
○ differs from adult
○ occupies separate ecological niche
○ Indirect dev.- includes larval stage
○ biphasic life cycle- aqu. invertebrates, bottom dwelling (benthic)
adult & floating (planktonic)/swimming larva
○ larva- anatomically diff. from juvenile & adult
○ must locate & settle in a habitat suitable for adult
○ juvenile- immature form of adult; this transform.= metamorphosis
○ metamor.- ranges from gradual addition of new str. & slight
rearrangement of tissues to a juvenile from undifferentiated
embryonic reserve cells in larval body
○ process by which juvenile differentiates from larval embryonic
reserve cells resembles budding

○ larvae of aqu. inverte. maybe benthic or planktonic

○ larval period- short or long
○ lecithotrophs (=yolk-feeders) - short-lived larvae commonly feed on
maternal supply of yolk that carries over the egg into embryonic
endoderm
 ○ planktotrophs- most long-lived larva initially lecithotrophic then a
functional gut differentiates, then sustains by feeding on plankton
● Indirect developers :
○ whose life cycle included planktonic larval stage produces many
microlecithal eggs
■ as they feed in diff. habitat
■ not compete with benthic adults for food
■ if long-lived, disperse widely
■ long planktonic period inc. risk of predation & death→ so large
no. of eggs produced
○ whose life cycle has planktonic lecithotrophic larval stage produces
fewer eggs than spe. of planktotrophic larvae
■ each egg- large & substantial amt. of yolk
○ short planktonic existence- reduces risk of predation, allows only
limited dispersal
● Direct development- bypass larval stage, embryo directly into juveniles
● primitive pattern- indirect life cycle with ext. fert. & planktonic dev.
● Reproductive adaptations- enhance ferti. & embryo survival
○ increasing synchrony (male & female gametes are produced &
released at same time)
○ proximity (gametes released near each other)
○ Synchronous production and gamete release triggered largely by
environ. signals- temp., light, & tides
○ Hermaphroditism= presence of both male & female gonads in the
same individual; common adaptation
■ low pop. density
■ adults live permanently attached to surfaces
■ assures nearby individual is potential mate
■ Most herm. animals cross-fertilize rather than self-fertilize
● selfing limits genetic variation
● enhances expression of deleterious recessive alleles
○ Gonochorism (diocy)- occurance of separate male & female
individuals; opp. of herm.
● Adaptations-
○ survival of offspring
○ maternal provisioning of nutrients & physical protection of eggs &
embryos- most imp.
● Oviparous spe. that prod. eggs before or immediately after fert.provide
shortest period of paternal protection
 ● Viviparous spe. have int. fert., retain (gestate) embryos in maternal body,
release differentiated offspring- larval or juvenile
○ nutrients from-
■ matrotrophically- directly from mother via placenta
■ lecithotrophically- from yolk stored in egg
○ Brooding- when eggs are released from mother but then retained
into her body

FUNCTIONAL CONSEQUENCES OF BODY SIZE:

● evolution of multicell. animals from unicell. protozoans- resulted in new traits:
○ tissue-level skeleton
○ larger body size of metazoans
● size- less than 1 um to ms
○ prokaryotic org.- <1um to 2um
○ unicell. protozoans- 2um to 0.5 mm
○ multicellular ani.- 0.5 mm to >1m
● fund. changes in body design must have occurred at approx. 2um to 0.5mm
○ transition at 2um from prokaryotic to eukar. cells
○ another redesign at 0.5mm; marking the transition from protozoa to
metazoa
● Intro. to Eumetazoa- 1st animals to evolve muscle & nerve
● diff. body sizes of prokar., proto, metazoans
○ 1um bacterium- Pneumococcus; correspond to mustard seed
○ 100um protozoan- dinoflagellate; corr. grapefruit
○ 10cm metazoan- sea urchin; corr. Astrodome

SIZE & COMPARTMENTALIZATION-

● cell specialisation segregates funct.- similar to div. of labor
● each proficient at specific essential task
● improves- efficiency, work output-to-input ratio
● efficiency of cell specialisation is linked with the interdependence of those
spec., func. compartmentalised cells
○ their evolution relied on func. compart. of new cellular level & their
func. integration
● Funct. compart. & integration- not limited to metazoans
○ protozoans- compart. are intracellular organelles
○ div. of labor at organelles level & integration of their specialised
func. by intracellular chemical messengers→ greater size & activity
of eukar. cells
○ imp. themes in metazoans evolution
 ● metazoans inherited eukar. cell organelles but evolved a new level of
compart.- specialisation of cells within the multicellular body
○ skeleton support with tissues
○ funct. compart. includes organ & organ system
○ chemical environ. of compartment regulated by its borders
○ systems of control & integration- nervous, endocrine & circulatory
sys.
○ eg- when muscle tissue 1st evolved, nervous tissue accompanied it
to control its contraction
SIZE, S.A. & VOLUME-
● for an org. of const. shape, change in body size disproportionately
changes the body’s surf. area in relation to its vol.
● area-to-vol. ration is imp.
○ exchange of substances- gases, nutrients & wastes
○ metabolic demand of a body is related to its vol.
● a solid 3-D body becomes larger, surf. area inc. in proportion to r2, but vol.
inc. more rapidly in proportion to r3
○ reach a size at which it’s surf. area (supply) is inadequate to serve
its vol. (demand)
● size of prokar. bacterium, eukar. protozoan & metazoan- all cubical shape
○ for 1um bacterium, S.V. is 6:1
○ 10um protozoan is 0.6:1 (10 fold dec. in area for each unit of vol.)
○ 1cm metazoan is 0.006:1 (100-fold drop)
○ mammalian cell is 200000:1
● calculated dec. in S.V. for eukar. cells is more fiction than fact
○ because eukar. cells rarely retain smooth surfaces or perfect
geometric shapes
○ their surf folded or bear outgrowths (microvilli, pseudopodia) that
inc. area
○ eukar. animal cells with enhanced surfaces can supply their vol.
metabolic demand at a greater rate than can prokar. cells
○ organelles like Golgi & ER are surface ingrowths that inc. area for
metabolic machinery & amplify its metabolic capacity
● inherited surface-enhanced
○ earliest metazoans inh. eukar. cells but their larger body req.
improvement in area-to-vol. ratio
○ primitive metazoans achieved this by arranging their cells in 2 thin
2-D sheet over ECM
○ eg- sponges or earthworms arrange cell sheets in the form of hollow
tubes or tubes in tubes
 ○ eg- flatworms, roundworms, and slime molds, are flat and thin or
long and slender as a means of improving the area-to-volume ratio

SIZE & TRANSPORT-

● Diffusion- process by which molecules in a liquid or gas enter and leave
cells & tissues
 ○ results from random movement of heated (above absolute 0) molec.
of liquids & gas
○ motion is random; if O2 is in high conc. outside the cell, then more
O2 will enter the cell than inside → conc. diff. result in net movement
of O2
● diffusion is slow over greater distances
○ as time req. to achieve a conc. (C) is proportional to distance sq.
from source
○ For aquatic animals, the effective diffusion distance is roughly 0.5
mm → this means that a solid, metabolically active, cellular body will
be diffusion limited at a diameter of approx. 1 mm.
● some metazoans & protozoans rely on diffusion for int. & ext. transport
○ larger proto & metazoans have altered their body shapes to remain
within the limits of diffusion
■ becoming slender & thread-like or flat & thin
■ in both- body radius is small & S.A. is large in relation to vol.
○ eg-
■ Protozoans- large forams with numerous radiating thread-like
reticulopods;
■ giant multinucleated amebas are flat & thin;
■ even largest known cell ( a multinucleated ameba) is the slime
mold Physarum polycephalum area 5.54 m sq., 1mm
thickness (S.V= 1:1)
■ metazoan flatworms- common planaris - flat & thin as tissue
paper
■ metazoan worms whose cylindrical bodies are cellular from
surface to core are thin or thinner than sewing thread
● for bodies exceeding 1 mm in diameter → diffusion alone not sufficient
○ cells farthest from surf. may die
○ Circulatory system- an internal transport system that provides a
mass flow of fluid (= convention) & frees metazoans from body-size
& -shape limitations imposed by simple diffusion
■ animals- cilia, muscles both circulates the fluid
■ fluid transports metabolites b/w supplies & demands
● metazoan circulatory system may have evolved from internalised surface
tissue that formed circulatory vessels
○ similar to water-filled channels (aquiferous system); in sponges
○ or simple invagination of 1 body side; in Hydra & coelenteron
○ during gastrulation in the dev. of most metazoans
 ○ Hemal & coelomic systems- bilaterally symmetrical animals evolved
functionally specialised circulatory systems
○ Circulatory system was a precondition for the evolution of large
body size
SIZE & METABOLISM-
● generally, metabolic rate inc. with body size- bigger animals consume
more power per unit time than smaller animals but as body size inc. each
additional unit of mass req. less than 1 additional unit of power
○ eg- large metazoan composed of 10,000 cells consumes more
energy per unit time than a single celled protozoan does, but not
10,000 times more energy
or
metabolic rate of that single protozoan cell exceeds that of any 1 of
the 10000 metazoan cells
○ 1g of shrew tissue consumes more energy than 1g of elephant
tissue does
■ small animals have greater mass-specific power consumption
■ large animals might receive a mass-specific discount in power
consumption
○ offset
■ b/w the protozoans and metazoan poikilotherms represents
8-times-greater power consumption of a metazoan as
compared with a protozoan of equal mass;
■ b/w poikilotherm (cold-blooded) and homeotherm lines shows
29-times-greater power consumption of a homeotherm vs. a
poikilotherm of equal weight
■ both → represents the cost of maintaining a const. body temp.
○ Q. Why does it require 8 times more energy to power a metazoan
than a protozoan of equivalent size.
■ b/c of extracellular circulatory system; Metazoans may pay a
surcharge for the power needed to circulate fluid using cilia,
muscles, or both, & ultimately for the evolution of large body
size

● f
 ADVANTAGES OF LARGE BODY SIZE-
● Large body size of metazoans-
○ confers a mass-specific discount on metabolic rate & other adv.
○ lowered metazoan vulnerability to protozoan predation (chances are
less; eg- small metazoans- rotifers & gastrotrichs are eaten by
protozoans)
○ due to large body + tissue specialisation → efficient at eating
protozoans (imp. adv.)
○ eg- sponges can consume particulates including eukar. cells in the
submicro.-50 um size range [wide range- as their food filter not
restricted to collar cells alone; also includes tissue level
(multicellular) component- not available to protozoan] whereas
choanoflagellates are restricted to smaller ones
○ can move at greater speed too
● multicellularity buffers the individual against environ. damage
○ multicellularity & regenerative ability- enable body to tolerate cell
loss
■ eg- novel toxins ingested with food- damage some cells but
don’t kill the individual
ONTOGENY & PHYLOGENY:
● Ontogeny- developmental time course over which growth & differentiation
convert zygote to multicellular adult
○ has dev. stages; variations subject to natural selection- potential for
evolution of new forms
 ■ eg- life cycle of segmented worm that includes benthic adult &
planktonic larva;
● random variation→ sexually mature larvae but never grew to
adult form
● favored by natural selection; b/c exploited the novel environ. &
new spe. resembles the larvae not adult
● ⇒ sexual maturity appeared sooner
■ eg- toothed whales (dolphins, orcas, sperm whales)
● 1 incisor tooth grew either fastly or extended period res. in
enormous unicorn-like tusk
● a potent sexual signal, new whale spe. evolved
● ⇒ inc. in the rate/period of dev. of organ or str.
■ in both- varients arouse from a change in timing of developmental
events
● Heterochrony (=changed time)- changes in the timing of developmental events
○ at any stage; affect any component; permits radical (& moderate)
evolutionary change
○ 2 categories-
■ pedomorphosis (=restrained shaping)- if descendant spe.’s trait
resembles ancestral dev. trait and this is larval or juvenile in form
● smaller & anatomically simpler descendants
● short generation time
● eg- spe. in temporary pools, plankton community, symbiotic
spe.- parasitic barnacles (nonpedomorphic female modified to
absorb nutrients from host & produce myraid eggs & offspring
too infect new hosts; pedomorphic male is dwarf- little more
than sexually matured larva, parasitic on females, only role- to
provide sperms; tiny- not compete with female for nutrition
● heterochrony in a family of segmented worms (polychaetes)
enabled the small-bodied pedomorphic descendant to
colonize the minute water-filled spaces between sand grains
(an interstitial community)
■ peramorphosis (=extended shaping)- if descendant’s trait develops
to an extent beyond that found in ancestor; eg- narwhal’s enormous
tooth
● larger descendants & anatomically complex
● long generation times- favored in const. or predictable
environ.- deep sea, upwelling areas of sea, coral reefs
 ● in worm eg- morphogenesis of a functional gonad in larval stage of ancestral life
cycle- a case of pedomorphosis; but occurance of gonads at early stage- result
from [3 forms of pedomorphosis]
○ progenesis- normal (ancestral) rate of dev. early onset of sexual maturity,
arresting somatic dev.
○ neoteny- normal (ancestral) rate of dev. leading to sexual maturity with
dec. dev. rate of somatic str. i.e. differentiation of body lags permanently
behind reproductive dev.
○ postdisplacement- normal (ancestral) rate of dev. with delayed onset of
somatic str. resulting again in sexually mature juvenile body
● Ancestral ontology :- (ontology- nature of being and existence)
○ Progenesis: developmental rate is unchanged, but an early onset of
gonad differentiation arrests further body (somatic development, thus
shortening the developmental period).
○ Neoteny: the developmental period and onset are unchanged and the
rate of gonad differentiation also is unchanged, but the rate of somatic
differentiation is slowed.
○ Postdisplacement: the period and rate of development are unchanged,
and the onset of gonad differentiation also is unchanged, and the onset of
somatic differentiation is delayed.
○ Hypermorphosis: developmental rate and somatic onset are unchanged,
but the developmental period is extended and the onset of gonad
differentiation is delayed.
○ Acceleration: developmental period and onset are unchanged, but rate of
somatic differentiation is increased.
○ Predisplacement: developmental period and rate are unchanged, but
onset of differentiation is early.

EVOLUTIONARY ORIGINS :
ORIGIN OF METAZOA
● common ancestry with some unicell. org.
○ Colonial theory- metazoa is derived from colony of flagellated
protozoa
○ Syncytial theory- metazoans evolved from a multinucleate but
unicell. plasmodium similar to slime mold or a ciliate protozoan, later
memb. evolved to prod. cell boundary around each nuclei;
■ support from dev. of mold/ Drosophila insect in which
multinucleate stage followed by cellularization to form
multicell. body;
■ phylogenetic analysis contradicts it & favor colonial theory
● modern view- Colonial theory= premetazoan (a protozoan) consisted of a
small spherical colony bearing a surface layer of flagellated cells used for
locomotion & feeding
○ colony originated from cell (form by mitosis); daughter cells not
separate after cell div.
○ daug. cells held by a proteinaceous ECM (occupies much int.
space) & deeply embedded
○ choanoflagellates each bore a single collared flagellum
○ few nonflagellated cells, capable of giving rise to flagellated cells &
gametes scattered in subsurface ECM
● Flagellated, or monociliated, collar cells-cells with single flagellum & collar
or microvilli : widespread metazoans
● ciliated cells of lower metazoans- sponges, jellyfishes, sea anemones &
corals : monociliated
● Choanoflagellates & metazoans → sister taxa (data indicate)
● ECM- cells are attached & mode of movement - universal
● cellular specialisation - eggs, sperms : evolved in colonial flagellated eg-
Volvox
● Volvox-
○ analog for ancestral metazoan; shows how multicell. org. evolved
from unicell. ancestor
○ not the ancestor of metazoa
○ an autotrophic org., plant-like cells
○ multicellularity evolved 50-75 million yrs. ago; as metazoans- 600
million yrs.
○ volvocids evolved multicellularity in || with metazoans

○ colonie of Proterospongia haeckeli resembles premetazoan

 ■ consists flagellated collar cells in gel. ECM surf.
■ cells undergoing div. & lacks flagella- deep in ECM
● choanoflagellate collar cells are identical to collar cells (choanocytes) in
metazoan sponges
○ both- single flagellum; surrounded by collar of microvilli
○ flagellum anchored in cells by microtubules; radiate form flagellar basal
body
● 1st hypothetical metazoan= protometazoan
○ different form premetazoan [4 reasons]
■ surf. cell closely adjoined; intercellular communication; regular
barrier b/w ext. & ECM
■ close associated excluded ECM from b/w the cells; separate ECM
into ext. & int. layers- each independent func.
■ body polarised along ant.-post. axis
■ layer separation & body polarization promoted cell specialisation
ORIGIN OF POLARITY & CELL SPECIALIZATION :
● most motile Protozoans have
○ leading- anterior & trailing- posterior ends
○ if attached: oral free ends & aboral attached ends
● in Metazoans; polarization from eggs
○ during oogenesis, eggs express a rudimentary flagellum & collar of
microvilli at a site on the cell surf. corresponds to animal pole of
zygote
○ animal pole corr. to ant. end of larva (though post. in others)
○ => line of descent form polarity of a choanocyte to primary ant.-post.
of metazoan body
○ prometazoan polarised along ant.-post. axis
● environ. condn. for polarity evolution:
○ for aqu. metazoan → light, temp., oxygen & food availability
○ eg- if food avail. related to light (photosyn.), natural selection would
favor any individual variant, one capable of detecting & following
light gradient; directional-sensitive sensory capability coupled with a
directional (polarised) locomotary system
○ capable of tracking resource conc. gradient
○ biologist Lewis Wolpert- body polarity evolved from attachment to a
substratum
■ eg- attachment of rock in water places an org. at an interface
(steep gradient), once attached variants favor that adhere well
at the attached end & perform tasks- feeding at the opposite
end; leads to polarised body
● after polarity, creates Environ. gradient → for differential expression of
traits; enhanced:
○ memb. sensitivity to environ. stimuli favor at the ant. end- as 1st to
encounter changes
○ flagellar growth, density, or activity - favor at the equator or widest
part - these locations best to contribute to locomotion
○ division capacity to growth - favor at post. end - interfere least to
locomotion
○ motility along polar axis- promote cell. spec.- cells occupy diff. fixed
positions in environ. gradient
● Acc. to biologist Leo Buss = origin of metazoan cell specialisation related
to conflict b/w demands of growth & locomotion
○ Volvox, Proterospongia & planktonic blastula stages of metazoans
req. flagellated surf. for locomotion, but flag. cells cannot divide by
mitosis b/c centrioles needed to form mitotic spindle are already in
use as flagellar basal bodies
■ metazoan flag. cell only divide after flagellum regresses & its
basal bodies freed
● growth in premetazoan composed solely of flag. cells are
○ 1- enlargement of existing cells (Volvox & few postembryonic
micrometazoans grows by cell enlarg. only)
○ 2- disassembly of flagella, cell div., & then flagellar reassembly
(regression of flagella compromises locomotion)
○ 3- div. of scattered cells in embryo, while others retain flagells
○ 4- div. of localised cells(mitotically active) set aside for growth
[preferred]; evolutionary event- setting aside germ cells capable of
meiosis
● Surface flagella- necessary for locomotion in protometazoan
● optimal position for set-aside growth cells:
○ Buss- at surf. result in outgrowth of locomotory cells/ tumorlike
appendages = -vely impact motility; internalising growth cells-
neither distort the surf. nor interfere with locomotory cells
○ Protometazoan did internalise its growth cells & process=
gastrulation
○ internalisation of cells capable of mitosis- in Volvox (flag. surf. cells
& invaginated subsurf. pockets of non-flag. dividing cells {gonidia-
give rise to either new colonies or germ cells}) , P. haeckeli (it’s
colony- by addn. of new surf. choanocytes that originates by mitosis
in subsurf. gel)
● 2 layers of differentiated cells in early metazoans (req. for locomotion &
growth); cell layering established
○ tissues
○ ext.-int. gradient
● surf. cells- direct access to gases & raw mat.; constrained for locomotion
& ext. environ.
● interior cells- became specialised for repro., nutrient storage & food
digestion (by gut evolution)
ORIGIN OF COMPLEXITY :
● metazoans evolved from protozoans; spec. for diff. func.
● Metazoa= as replication of similar units (cells) followed by unit
specialisation & integration into organism at a new & higher level of
complexity → Replication-specialisation-integration of units (a general
pattern)
● ciliates- replication, specialisation & integration of cilia → largest, most
diverse, active & complex protozoan cells
● metazoans → body composed of series of similar segments eg- in
earthworms, crustaceans; later became specialised & integrated into
head, thorax & abdomen