The Evolutionary

History of
Biological Diversity
Bui Hong Thuy, Ph.D.
School of Biotechnology,
International University
Email: bhthuy@hcmiu.edu.vn

1
 OUTLINES
1. Plant Diversity I: How
Plants Colonized Land

2. Plant Diversity II: The

Evolution of Seed Plants

3. An Introduction to
Animal Diversity

4. Animal Development

2
 OUTLINES
1. Plant Diversity I: How
Plants Colonized Land

2. Plant Diversity II: The

Evolution of Seed Plants

3. An Introduction to
Animal Diversity

4. Animal Development

3
 1.1. Land plants evolved from
green algae

 Since colonizing land at least 475 million years

ago, plants have diversified into roughly
290,000 living species.
 Green algae called charophytes are the closest
relatives of land plants.
 Note that land plants are not descended from
modern charophytes, but share a common
ancestor with modern charophytes.

4
Morphological and Molecular
Evidence
Land plants share key traits only with green
algae charophytes:
• DNA comparisons of both nuclear and chloroplast
genes.
• Rose-shaped complexes for cellulose synthesis.
• Peroxisome enzymes - minimize loss from
photorespiration.
• Structure of flagellated sperm.
• Formation of a phragmoplast - allignment of
cytoskeletal elements and Golgi vesicles for cell plate.

5
Adaptations Enabling the Move to Land

 In green algae charophytes a layer of a durable 
polymer called sporopollenin prevents 
dehydration of exposed zygotes.
 The movement onto land by charophyte 
ancestors provided advantages:  unfiltered sun, 
more plentiful CO2, nutrient‐rich soil, and few 
herbivores or pathogens.
 Land presented challenges: a scarcity of water 
and lack of structural support.

6
 Three Clades are candidates for Plant Kingdom

Red algae

ANCESTRAL

Viridiplantae
ALGA Chlorophytes

Streptophyta
Charophytes

Plantae
Embryophytes

7
 Alternation of Generations and
Multicellular Dependent Embryos
 The multicellular gametophyte is haploid and
produces haploid gametes by mitosis.
 Fusion of the gametes gives rise to the diploid
sporophyte, which produces haploid spores by
meiosis.
 The diploid embryo is retained within the tissue
of the female gametophyte. Nutrients are
transferred from parent to embryo through
placental transfer cells.
 Land plants are called embryophytes because
of the dependency of the embryo on the parent.

8
Haploid (n) Gamete from
Gametophyte (n)
phase another plant

Mitosis Mitosis n
n
n
n Spore
Gamete

Meiosis Fertilization

2n Zygote

Diploid (2n)
phase Alternation of generations =
Sporophyte (2n) Derived traits of land plants 9
Derived Traits of Land Plants
Multicellular Dependent Embryos

Embryo
2 µm Maternal tissue

Wall ingrowths 10 µm
Placental transfer cell
(outlined in blue)
Embryo (LM) and placental transfer cell (TEM)
of Marchantia (a liverwort) 10
Walled Spores Produced in Sporangia

 The sporophyte produces spores in

organs called sporangia.
 Diploid cells called sporocytes undergo
meiosis to generate haploid spores.
 Spore walls contain sporopollenin, which
protects against dessication making them
resistant to harsh environments.

11
Derived Traits of Land Plants:
Walled Spores Produced in Sporangia

12
 Multicellular Gametangia
Gametes are produced within ‘sex
organs’ called gametangia.
Female gametangia, called
archegonia, produce eggs and are
the site of fertilization.
Male gametangia, called antheridia,
are the site of sperm production and
release.

13
 Derived Traits of Land Plants:
Multicellular Gametangia - ‘sex organs’
Archegonium
Female gametophyte with egg

Antheridium
with sperm

Male
gametophyte

Archegonia and Antheridia of Marchantia (a liverwort) 14

 Apical Meristems

 Apical meristems are growth regions at

plant tips, allowing plants to sustain
continual growth in their length.
 Cells from the apical meristems differentiate
into various tissues.

15
 Apical Meristems - Allow for Growth in Length
throughout Plant’s Lifetime.
Apical Developing Apical meristems
meristem leaves
of shoot

Apical meristem
Shoot 100 µm of root Root 100 µm

Derived Traits of Land Plants 16

 A Vast Diversity of
Modern Plants
 Ancestral species gave rise to land plants
which can be informally grouped based
on the presence or absence of vascular
tissue.
 Nonvascular plants are commonly called
bryophytes.
 Most plants have vascular tissue; these
constitute the vascular plants:
seedless vascular and seed plants.

17
 Seedless vascular plants can be divided into
clades:
– Lycophytes (club mosses and their relatives)
– Pterophytes (ferns and their relatives).

 A seed is an embryo and nutrients surrounded

by a protective coat.
 Seed plants form a clade and can be divided into
further clades:
– Gymnosperms, the “naked seed” plants including the
conifers / cone = sex organ
– Angiosperms, the flowering plants including monocots
and dicots / flower = sex organ

18
 Highlights of Plant Evolution
1 Origin of land plants (about 475 mya)
2 Origin of vascular plants (about 420 mya)
3 Origin of extant seed plants (about 305 mya)
Liverworts

(bryophytes)
plants
Nonvascular

Land plants
ANCES- 1 Hornworts
TRAL
GREEN
ALGA Mosses

Lycophytes (club mosses,

Vascular plants
plants
vascular Seed plants
Seedless
spike mosses, quillworts)

2 Pterophytes (ferns,
horsetails, whisk ferns)

Gymnosperms

3
Angiosperms

500 450 400 350 300 50 0

Millions of years ago (mya)

Several hypotheses about relationships between plant groups are actively under
debate. The dotted lines indicate groups whose evolutionary relationships are unclear
19
1.2. NonVascular plants have life cycles
dominated by gametophytes

 Bryophytes are nonvascular and represented

today by three phyla of small herbaceous
(nonwoody) plants:
– Liverworts, phylum Hepatophyta
– Hornworts, phylum Anthocerophyta
– Mosses, phylum Bryophyta
 Mosses are most closely related to vascular plants.

 Gametophytes are dominant: larger and longer-

living than sporophytes. Sporophytes are present
only part of the time and dependent on the
gametophytes.
20
Life Cycle of a Bryophyte Raindrop
> Moss
Gametophyte is the Sperm Mature gametophytes
Dominant Generation produce flagellated
“Bud” Antheridia sperm in antheridia
Key
Male and an egg in each
gametophyte archegonium
Haploid (n) (n)
Diploid (2n) Protonema
(n) “Bud”
Egg
Sperm swim
Spores Gametophore
Archegonia through a film of
Spore Female water to reach
dispersal gametophyte (n)
and fertilize the
Peristome Rhizoid egg

Sporangium FERTILIZATION
MEIOSIS Seta (within archegonium)
Capsule Zygote
Mature (2n)
sporophytes (sporangium)
Foot Embryo

Archegonium
Young
sporophyte
2 mm

(2n)
Capsule with Female 21
peristome (SEM) gametophytes
Bryophyte Structures

Gametophore of
Thallus female gametophyte

Sporophyte

Foot
Seta

Capsule
Marchantia polymorpha, (sporangium)
a “thalloid” liverwort

500 µm
Marchantia sporophyte (LM)
22
The Ecological and Economic
Importance of Mosses
Bryophytes / Moss may help retain Nitrogen in the
soil, RESULTS
an Ecological Advantage
6
Annual nitrogen loss

5
4
(kg/ha)

3
2
1
0
With moss Without moss 23
Sphagnum,
or peat moss

Economic and
archaeological
significance

(a) Peat being harvested from a peat bog.

(b) “Tollund Man,” a bog mummy: The acidic, oxygen

poor conditions can preserve bodies.
24
1.3: Ferns and other seedless vascular
plants were the first plants to grow tall
 Bryophytes and bryophyte-like plants were the
vegetation during the first 100 million years of plant
evolution.
 Vascular tissue allowed vascular plants to grow tall.
 Seedless vascular plants have flagellated sperm
and are usually restricted to moist environments.
 Vascular plants are characterized by:
• Life cycles with dominant sporophytes
• Vascular tissues called xylem and phloem.
• Well-developed / true roots and leaves.

25
 Life Cycle of a Seedless Vascular Plant -
Fern Dominant Sporophyte
Key
Haploid (n)
Diploid (2n)
Spore Young Antheridium
Spore (n) gametophyte
MEIOSIS dispersal

Sporangium Mature Sperm

gametophyte
(n) Archegonium
Egg
Mature
Sporangium sporophyte New Zygote
(2n) sporophyte (2n) FERTILIZATION
Sorus

Gametophyte

Fiddlehead
26
 Evolution of Roots and Leaves
 Roots are organs that anchor vascular plants and
enable plants to absorb water and nutrients from
the soil.
 Leaves are organs that increase the surface area
of vascular plants for capturing more solar energy
used for photosynthesis.
Hypotheses for Evolution of Leaves
Overtopping
growth Megaphyll
Vascular tissue Sporangia Microphyll

Other stems Webbing

become re- develops.
duced and
flattened.

(a) Microphylls - single veined leaves (b) Megaphylls - branching leaf veins
27
Homosporous spore production
Typically a Eggs
Sporangium Single bisexual
on sporophyll type of spore gametophyte
Sperm

Heterosporous spore production

Megasporangium Female
on megasporophyll Megaspore gametophyte
Eggs

Microsporangium Male
on microsporophyll Microspore Sperm
gametophyte

28
 Seedless Vascular Plants
Lycophytes (Phylum Lycophyta)
2.5 cm

Isoetes Strobili
gunnii, (clusters of
a quillwort sporophylls)
Selaginella apoda,
a spike moss

1 cm

Diphasiastrum tristachyum, a club moss

29
 The Significance of Seedless
Vascular Plants
 Increased photosynthesis may have helped produce
the global cooling at the end of the Carboniferous
period.
 The decaying plants of these Carboniferous forests
eventually became coal = fossil fuel.

Artist’s depiction of a
Carboniferous forest
based on fossil evidence
30
Derived Apical meristem Developing
of shoot leaves
Traits of Gametophyte
Mitosis
Mitosis
Plants n n

n n
Spore Gamete

MEIOSIS FERTILIZATION

2n Zygote

Mitosis
Haploid
Sporophyte
Diploid
1 Alternation of generations 2 Apical meristems

Archegonium Antheridium Sporangium Spores

with egg with sperm

3 Multicellular gametangia 4 Walled spores in sporangia 31

 OUTLINES
1. Plant Diversity I: How
Plants Colonized Land

2. Plant Diversity II: The

Evolution of Seed Plants

3. An Introduction to
Animal Diversity

4. Animal Development

32
2.1. Seeds and pollen grains are key
adaptations for life on land
 In addition to seeds, the following are common to
all seed plants:
– Reduced gametophytes
– Heterospory
– Ovules
– Pollen

What human reproductive

organ is functionally
similar to this seed?

 A seed consists of an embryo

and nutrients surrounded by a
protective coat.
33
 Gametophyte / sporophyte relationships
in different plant groups
PLANT GROUP
Mosses and other Ferns and other Seed plants
seedless (gymnosperms and angiosperms)
nonvascular plants vascular plants
Reduced, independent Reduced (usually microscopic),
Gametophyte Dominant (photosynthetic and dependent on surrounding
free-living) sporophyte tissue for nutrition
Reduced, dependent
Sporophyte on gametophyte Dominant Dominant
for nutrition
Gymnosperm Angiosperm
Sporophyte Microscopic female
(2n) gametophytes (n) inside
ovulate cone
Microscopic
Sporophyte
female
(2n)
gametophytes
(n) inside
Gametophyte
these parts
(n)
of flowers
Example
Microscopic
male
Microscopic male gametophytes
gametophytes (n) (n) inside
inside pollen these parts
cone of flowers
Sporophyte (2n) Sporophyte (2n)
Gametophyte
(n)
34
34
 Heterospory:
The Rule Among Seed Plants

 The ancestors of seed plants were likely

homosporous, while seed plants are heterosporous
 Megasporangia produce megaspores that give rise
to female gametophytes.
 Microsporangia produce microspores that give rise
to male gametophytes.
• Gymnosperm megaspores have one integument.
• Angiosperm megaspores usually have two
integuments.

35
 Ovules and Production of Eggs
• An ovule consists of a megasporangium, megaspore,
and one or more protective integuments.
• A fertilized ovule becomes a seed.
From ovule to seed
in a gymnosperm
Integument

Spore wall

Immature
female cone

Megasporangium
(2n)

Megaspore (n)

(a) Unfertilized ovule 36

 Pollen and Production of Sperm
 Microspores develop into pollen grains, which
contain the male gametophytes.
 Pollination is the transfer of pollen from the male
to the female part containing the ovules.

From ovule to seed

Female
in a gymnosperm gametophyte (n)
Spore wall
Egg nucleus (n)

Male gametophyte
(within a germinated
Discharged
pollen grain) (n)
sperm nucleus (n)

Micropyle Pollen grain (n)

(b) Fertilized ovule 37

The Evolutionary Advantage of Seeds
 Seeds provide some evolutionary advantages over
spores:
– They may remain dormant for days to years, until
conditions are favorable for germination.
– They may be transported long distances by wind or
animals.
Seed coat
From ovule to seed (derived from
integument)
in a gymnosperm

Food supply
(female
gametophyte
tissue) (n)

Embryo (2n)
(c) Gymnosperm seed (new sporophyte) 38
 Gymnosperms bear “naked”
seeds, typically on cones
 The gymnosperms have “naked” seeds not
enclosed by ovaries and exposed on modified
leaves - cones. There are four phyla:
– Cycadophyta (cycads)
– Gingkophyta (one living species: Ginkgo biloba)
– Gnetophyta (three genera: Gnetum, Ephedra,
Welwitschia)
– Coniferophyta (conifers, such as pine, fir, and redwood).

39
 Seed plants can be divided into two
clades: gymnosperms and angiosperms.
 Gymnosperms were better suited than
nonvascular plants to drier conditions.
 Today, cone-bearing gymnosperms called
conifers dominate in the northern
latitudes.

40
 The Life Cycle of a Pine:
A Closer Look
• Three key features of the gymnosperm life cycle 
are:
– Dominance of the sporophyte generation.
– The transfer of sperm to ovules by pollen.
– Development of seeds from fertilized ovules.

• The life cycle of a pine provides an example.

41
Life Cycle
of a Pine Key
Haploid (n)
Ovule
Diploid (2n)

Ovulate Megasporocyte (2n)

cone
Integument
Pollen
cone
Microsporocytes
Mature (2n)
Megasporangium
sporophyte Pollen (2n)
(2n) Pollen grain
grains (n) MEIOSIS
MEIOSIS
Microsporangia
Microsporangium (2n) Surviving
Seedling megaspore (n)

Archegonium

Seeds Female
gametophyte

Food Sperm
reserves nucleus (n)
(n)
Seed coat
(2n)
Pollen
tube
Embryo
(2n)
FERTILIZATION Egg nucleus (n) 42
 The reproductive adaptations of
angiosperms include flowers and fruits
• Angiosperms are seed plants with reproductive
structures called flowers and fruits.
• They are the most widespread and diverse of
all plants.
• All angiosperms are classified in a single
phylum: Anthophyta.
• The name comes from the Greek anthos,
flower.

43
 Flowers - Specialized for Sexual Reproduction

 The flower is an angiosperm structure specialized

for sexual reproduction. It is a specialized shoot
with up to four types of modified leaves:
– Sepals - enclose the flower
– Petals - brightly colored and attract pollinators
– Stamens - produce pollen on their terminal anthers
– Carpels - consist of an ovary containing ovules at the
base and a style holding up a stigma, where pollen is
received.

44
Structure of an Idealized Flower
Stigma
Carpel
Stamen Anther
Style
Filament
Ovary

Petal
Sepal

Ovule

45
 Fruits
• A fruit typically consists of a mature ovary
but can also include other flower parts.
• Fruits protect seeds and aid in seed
dispersal.
• Mature fruits can be either fleshy or dry.
• Various fruit adaptations help disperse
seeds by wind, water, or animals to new
locations.

46
Fruits
Tomato
Ruby grapefruit

Nectarine

Hazelnut

Milkweed
47
 The Angiosperm Life Cycle
• The flower of the sporophyte is composed of both 
male and female structures.
• Male gametophytes are contained within pollen
grains produced by the microsporangia of anthers.
• The female gametophyte  = embryo sac, develops 
within an ovule contained within an ovary at the 
base of a stigma.
• Most flowers have mechanisms to ensure cross‐
pollination between flowers from different plants 
of the same species.

48
• A pollen grain that has landed on a stigma 
germinates and the pollen tube of the male 
gametophyte grows down to the ovary.
• Sperm enter the ovule through a pore opening
called the micropyle.
• Double fertilization occurs when the pollen tube 
discharges two sperm into the female gametophyte 
within an ovule.

49
Double Fertilization: Produces
Zygote 2n and endosperm (food) 3n
• One sperm fertilizes the egg forming a zygote.
• The other sperm combines with two nuclei and 
initiates development of food‐storing endosperm.
• The endosperm nourishes the developing embryo.
• Within a seed, the embryo consists of a root and 
two seed leaves called cotyledons.

50
 Key
Life Cycle of an Angiosperm
Haploid (n)
Diploid (2n) Microsporangium
Anther
Mature flower on Microsporocytes (2n)
sporophyte plant
(2n) MEIOSIS

Generative cell
Ovule (2n) Microspore Tube cell
(n)
Male gametophyte
Ovary (in pollen grain) Pollen
Germinating MEIOSIS (n) grains
Stigma
seed Megasporangium
(2n) Pollen
tube
Embryo (2n)
Endosperm (3n) Sperm
Seed coat (2n)
Seed Megaspore
(n)
Style
Antipodal cells
Female gametophyte Central cell
(embryo sac) Pollen
Synergids tube
Egg (n) Sperm
Nucleus of
developing (n)
endosperm
FERTILIZATION
(3n)
Zygote (2n)
Egg
nucleus (n) Discharged sperm nuclei (n) 51
 Angiosperm Phylogeny
• The ancestors of angiosperms and
gymnosperms diverged about 305 million
years ago.
• Angiosperms may be closely related to
Bennettitales, extinct seed plants with
flowerlike structures.
• Amborella and water lilies are likely
descended from two of the most ancient
angiosperm lineages.

52
 Angiosperm evolutionary history

Living
gymnosperms
Microsporangia Bennettitales
(contain
microspores) Amborella

Water lilies

Most recent common ancestor Star anise and

of all living angiosperms relatives
Monocots

Magnoliids

Eudicots

Ovules 300 250 200 150 100 50 0

Millions of years ago

(a) A possible ancestor of the (b) Angiosperm phylogeny

angiosperms?

53
 Angiosperm Diversity
 The two main groups of angiosperms are:
monocots - one cotyledon
eudicots (“true” dicots) - two cotyledons.
• More than one-quarter of angiosperm
species are monocots.
• More than two-thirds of angiosperm species
are eudicots.

54
Angiosperms Monocot
Characteristics
Eudicot
Characteristics

Embryos

Monocots One cotyledon Two cotyledons

and Leaf
venation

Eudicots
Veins usually Veins usually
parallel netlike

Stems

Vascular tissue
Vascular tissue usually arranged
scattered in ring

Roots

Root system Taproot (main root)

usually fibrous usually present
(no main root)

Pollen

Pollen grain with Pollen grain with

one opening three openings

Flowers

Floral organs Floral organs usually

usually in in multiples of
multiples of three four or five 55
 Evolutionary Links Between
Angiosperms and Animals
• Pollination of flowers and transport of seeds by
animals are two important relationships in
terrestrial ecosystems.
• Clades with bilaterally symmetrical flowers have
more species than those with radially
symmetrical flowers.
• This is likely because bilateral symmetry affects
the movement of pollinators and reduces gene
flow in diverging populations.

56
 Human welfare depends greatly
on seed plants
 No group of plants is more important to human
survival than seed plants.
 Plants are key sources of food, fuel, wood
products, and medicine.
 Our reliance on seed plants makes preservation
of plant diversity critical.

57
 Five Derived Traits of Seed Plants
Summary
Reduced Microscopic male and Male
gametophytes female gametophytes gametophyte
(n) are nourished and
protected by the Female
sporophyte (2n) gametophyte

Heterospory Microspore (gives rise to

a male gametophyte)

Megaspore (gives rise to

a female gametophyte)

Ovules Integument (2n)

Ovule
Megaspore (2n)
(gymnosperm)
Megasporangium (2n)

Pollen Pollen grains make water

unnecessary for fertilization

Seeds Seeds: survive

Integument
better than
unprotected
Food supply
spores, can be
transported Embryo
long distances
58
 OUTLINES
1. Plant Diversity I: How
Plants Colonized Land

2. Plant Diversity II: The

Evolution of Seed Plants

3. An Introduction to
Animal Diversity

4. Animal Development

59
 Animals Overview
• Animals are multicellular, heterotrophic eukaryotes
with tissues that develop from embryonic layers.
• There are exceptions to nearly every criterion for
distinguishing animals from other life-forms.
• 1.3 million living species of animals have been
identified.

60
 Animal Structure and
Specialization
• Nutritional Mode: Animals are heterotrophs
that ingest their food.
• Animals are multicellular eukaryotes.
• Their cells lack cell walls.
• Their bodies are held together by structural
proteins such as collagen.
• Nervous tissue and muscle tissue are unique
to animals.

61
 Reproduction and Development
 Most animals reproduce sexually, with the
diploid stage usually dominating the life cycle.
 After fertilization, the zygote undergoes rapid
cell division called cleavage.
 Cleavage leads to formation of a blastula.
 The blastula undergoes gastrulation, forming a
gastrula with different layers of embryonic
tissues.

62
 Animal Early Embryonic Development

Blastocoel
Cleavage
Endoderm
Blastula
Ectoderm

Archenteron
Zygote Eight-cell stage
Gastrulation Gastrula
Blastocoel Blastopore
Cross section
of blastula

63
• Many animals have at least one larval stage.
• A larva is sexually immature and morphologically 
distinct from the adult; it eventually undergoes 
metamorphosis.
• All animals, and only animals, have Hox genes
that regulate the development of body form.
• Although the Hox family of genes has been highly 
conserved, it can produce a wide diversity of 
animal morphology.

64
 The history of animals spans
more than half a billion years
• The animal kingdom includes a great diversity of 
living species and an even greater diversity of 
extinct ones.
• The common ancestor of living animals may have 
lived between 675 and 875 million years ago.
• This ancestor may have resembled modern 
choanoflagellates, protists that are the closest 
living relatives of animals.

65
 Body Plan -- Symmetry
 Animals can be categorized according to the
symmetry of their bodies, or lack of it.
 Some animals have radial symmetry.
 Two-sided symmetry is called bilateral symmetry.
 Animals with bilateral symmetry have:
– A dorsal (top) side and a ventral (bottom) side
– A right and left side
– Anterior (head) and posterior (tail) ends
– Cephalization, the development of a head. (Brain…)

66
 Animal Body Symmetry

(a) Radial symmetry

(b) Bilateral symmetry 67

 Body Plan -- Tissues
 Animal body plans also vary according to the
organization of the animal’s tissues.
 Tissues are collections of specialized cells
isolated from other tissues by membranous
layers.
 During development, three germ layers give
rise to the tissues and organs of the animal
embryo.

68
 Embryonic Germ Layers
 Ectoderm is the germ layer covering the
embryo’s surface.
 Endoderm is the innermost germ layer and
lines the developing digestive tube, called the
archenteron.
 Diploblastic animals have ectoderm and
endoderm.
 Triploblastic animals also have a middle
mesoderm layer; these include all bilaterians.

69
 Body Cavities
.
• A true body cavity is called a coelom and is
derived from mesoderm. Coelomates are
animals that possess a true coelom.
• A pseudocoelom is a body cavity derived from the
mesoderm and endoderm. Triploblastic animals
that possess a pseudocoelom are called
pseudocoelomates.
• Triploblastic animals that lack a body cavity are
called acoelomates.

70
 Coelom
Triploblastic Body covering
Animals Body (from ectoderm)

Cavities Tissue layer lining coelom

and suspending internal organs
Digestive tract (from mesoderm)
(from endoderm)

(a) Coelomate - true body cavity

Body covering
(from ectoderm)

Pseudocoelom Muscle layer

(from
mesoderm)
Digestive tract
(from endoderm)

(b) Pseudocoelomate
Body covering
(from ectoderm) Tissue-
filled region
(from
mesoderm)

Wall of digestive cavity

(from endoderm)

(c) Acoelomate - lack a body cavity 71

 Cleavage:
• In protostome development, cleavage is spiral 
and determinate.
• In deuterostome development, cleavage is radial 
and indeterminate.
• With indeterminate cleavage, each cell in the 
early stages of cleavage retains the capacity to 
develop into a complete embryo.
• Indeterminate cleavage makes possible identical 
twins, and embryonic stem cells.

72
 Protostome Deuterostome
Development Development
molluscs, annelids echinoderm, chordates

Eight-cell stage Eight-cell stage (a) Cleavage

Spiral and determinate Radial and indeterminate

Key
Coelom
Ectoderm
(b) Coelom formation
Mesoderm
Archenteron
Endoderm

Coelom
Mesoderm Blastopore Blastopore Mesoderm
Solid masses of mesoderm Folds of archenteron
split and form coelom. form coelom.

Anus Mouth (c) Fate of the blastopore

Digestive tube

Mouth Anus
Mouth develops from blastopore. Anus develops from blastopore. 73
 New views of animal phylogeny are
emerging from molecular data
• Zoologists recognize about three dozen animal
phyla.
• Current debate in animal systematics has led to
the development of two phylogenetic hypotheses,
but others exist as well.
• One hypothesis of animal phylogeny is based
mainly on morphological and developmental
comparisons.
• Another hypothesis is based mainly on molecular
data.

74
 “Porifera”

Metazoa
Cnidaria
ANCESTRAL
COLONIAL

Eumetazoa
Ctenophora
FLAGELLATE

Ectoprocta

Deuterostomia
Brachiopoda

Echinodermata

Bilateria
Chordata

Platyhelminthes

Protostomia Rotifera

A view of animal phylogeny Mollusca

based mainly on morphological Annelida

and developmental
comparisons Arthropoda

Nematoda 75
 “Porifera”
Silicea

Metazoa
Calcarea
ANCESTRAL
COLONIAL
FLAGELLATE Ctenophora

Eumetazoa
Cnidaria

Acoela

Deuterostomia
Echinodermata

Bilateria
Chordata

Platyhelminthes

Rotifera

Lophotrochozoa
Ectoprocta

Brachiopoda
A view of animal Mollusca
phylogeny based
Annelida
mainly on molecular
Ecdysozoa

data Nematoda

Arthropoda
76
 Points of Agreement
• All animals share a common ancestor.
• Sponges are basal animals.
• Eumetazoa is a clade of animals ‐ eumetazoans
with true tissues.
• Most animal phyla belong to the clade Bilateria, 
and are called bilaterians.
• Chordates and some other phyla belong to the 
clade Deuterostomia.

77
Common ancestor Animal Phylogeny
of all animals

Metazoa
Sponges
(basal animals)
Ctenophora

Eumetazoa
Cnidaria

True

Bilateria (most animals)

Acoela (basal
tissues bilaterians)
Deuterostomia

Bilateral
summetry Lophotrochozoa
Three germ
layers Ecdysozoa
78
 OUTLINES
1. Plant Diversity I: How
Plants Colonized Land

2. Plant Diversity II: The

Evolution of Seed Plants

3. An Introduction to
Animal Diversity

4. Animal Development

79
 Overview: A Body-Building Plan
 It is difficult to imagine that each of us began life as 
a single cell (fertilized egg) called a zygote.

80
1 mm
After fertilization, embryonic development proceeds
through cleavage, gastrulation, and organogenesis

 Important events regulating development

occur during fertilization and the three
stages that build the animal’s body
– Cleavage: cell division creates a hollow ball of cells
called a blastula
– Gastrulation: cells are rearranged into a three-
layered gastrula
– Organogenesis: the three germ layers interact and
move to give rise to organs.

81
• Fertilization is followed by cleavage, a
period of rapid cell division without growth.
• Cleavage partitions the cytoplasm of one
large cell into many smaller cells called
blastomeres.
• The blastula is a ball of cells with a fluid-
filled cavity called a blastocoel.

82
 0.25 mm 0.25 mm

Animal pole Blastocoel

Vegetal
Zygote 2-cell 4-cell 8-cell pole: Blastula
stage stage stage (cross
forming forming section)

83
 Gastrulation
• Gastrulation rearranges the cells of a
blastula into a three-layered embryo, called
a gastrula, which has a primitive gut.
• The three layers produced by gastrulation
are called embryonic germ layers:
– The ectoderm forms the outer layer
– The endoderm lines the digestive tract
– The mesoderm partly fills the space
between the endoderm and ectoderm.

84
 Gastrulation in the sea urchin embryo:
 The blastula consists of a single layer of cells
surrounding the blastocoel.
 Mesenchyme cells migrate from the vegetal pole
into the blastocoel.
 The vegetal plate forms from the remaining cells of
the vegetal pole and buckles inward through
invagination.
 The newly formed cavity is called the
archenteron.
 This opens through the blastopore, which will
become the anus.

85
 Gastrulation in a sea urchin embryo
Key

Future ectoderm
Future mesoderm
Future endoderm

Vegetal Pole Blastocoel

Filopodia
Animal Invagination pulling
pole
Blastocoel archenteron Archenteron - cavity
Blastocoel tip Blastopore
Mesenchyme
cells
Ectoderm

Vegetal Vegetal
plate pole Mouth
Mesenchyme
cells Mesenchyme Digestive tube
(mesoderm (endoderm)
Blastopore 50 µm forms future
skeleton)

Anus (from
blastopore)

86
 Gastrulation in the frog
 The frog blastula is many cell layers thick.
Cells of the dorsal lip originate in the gray
crescent and invaginate to create the
archenteron.
 Cells continue to move from the embryo surface
into the embryo by involution. These cells
become the endoderm and mesoderm.
– The blastopore encircles a yolk plug when
gastrulation is completed.
– The surface of the embryo is now ectoderm, the
innermost layer is endoderm, and the middle layer is
mesoderm.

87
Gastrulation SURFACE VIEW CROSS SECTION

in a frog Animal pole

embryo
Blastocoel

Dorsal lip
of blasto-
Dorsal lip
pore
of blastopore
Blastopore
Early
Vegetal pole
gastrula

Blastocoel
shrinking Archenteron

Ectoderm
Mesoderm
Blastocoel
remnant Endoderm

Archenteron
Key
Blastopore
Future ectoderm
Future mesoderm Late
gastrula Blastopore Yolk plug
Future endoderm 88
 Gastrulation in the chick
• The embryo forms from a blastoderm and sits
on top of a large yolk mass.
• During gastrulation, the upper layer of the
blastoderm (epiblast) moves toward the midline
of the blastoderm and then into the embryo
toward the yolk.
• The midline thickens and is called the primitive
streak.
• The movement of different epiblast cells gives
rise to the endoderm, mesoderm, and
ectoderm.

89
Gastrulation Dorsal Fertilized egg
in a chick Anterior
Primitive
streak
embryo
Left Right Embryo

Yolk
Posterior
Ventral
Primitive streak

Epiblast

Future
ectoderm

Blastocoel
Migrating Endoderm
cells Hypoblast
(mesoderm) YOLK
90
 Organogenesis
• During organogenesis, various regions of the
germ layers develop into rudimentary organs.
• The frog is used as a model for
organogenesis.
• Early in vertebrate organogenesis, the
notochord forms from mesoderm, and the
neural plate forms from ectoderm.

91
 Early organogenesis in a frog embryo
Neural folds Eye Somites Tail bud

Neural Neural plate

fold

SEM
1 mm
1 mm
Neural tube Neural
Neural Neural crest
Notochord cells
fold plate
Neural crest Coelom Somite
cells

Notochord
Ectoderm
Archenteron
Mesoderm Outer layer (digestive
of ectoderm cavity)
Endoderm Neural crest (c) Somites
cells
Archenteron

(a) Neural plate formation

Neural tube

(b) Neural tube formation

92
 The neural plate soon curves inward, forming the
neural tube. The neural tube will become the
central nervous system = brain and spinal cord.
 Neural crest cells develop along the neural tube
of vertebrates and form various parts of the
embryo: nerves, parts of teeth, skull bones ...
 Mesoderm lateral to the notochord forms blocks
called somites.
 Lateral to the somites, the mesoderm splits
to form the coelom.

93
Organogenesis in a chick embryo is similar to that in a frog

Eye
Neural tube
Notochord Forebrain
Somite
Coelom Heart
Archenteron
Endoderm
Lateral fold
Mesoderm Blood
Ectoderm vessels

Somites
Yolk stalk
Yolk sac
These layers
form extraembryonic
membranes Neural tube
YOLK

(a) Early organogenesis (b) Late organogenesis

94
 ECTODERM MESODERM ENDODERM
Epidermis of skin and its Notochord Epithelial lining of
derivatives (including sweat Skeletal system digestive tract
glands, hair follicles) Muscular system Epithelial lining of
Epithelial lining of mouth Muscular layer of respiratory system
and anus stomach and intestine Lining of urethra, urinary
Cornea and lens of eye Excretory system bladder, and reproductive
Nervous system Circulatory and lymphatic system
Sensory receptors in systems Liver
epidermis Reproductive system Pancreas
Adrenal medulla (except germ cells) Thymus
Tooth enamel Dermis of skin Thyroid and parathyroid
Epithelium of pineal and Lining of body cavity glands
pituitary glands Adrenal cortex

95
 Human embryo development-
Fertilized egg Placenta
1 Cell Trophectoderm
(Zygote) (TE) cells

Inner cell
Blastocyst Mass (ICM)

Fetus

6,000,000,000 cells
(230 different
cell types)

96 96
 Blastocyst (Day 16)
Ectoderm
Mesoderm
Endoderm

Gastrulation

Nervous
Lines of
system,
digestive Muscle,
skin, lens
(Liver Bone, Blood,
of eye
Pancreas, Connective
stomach) & tissues
Respiratory
tract (đường
hô hấp :lung)

97
97
Ectoderm Endoderm Mesoderm
 Developmental Adaptations
of Amniotes
• Embryos of birds, other reptiles, and
mammals develop in a fluid-filled sac in a
shell or the uterus.
• Organisms with these adaptations are called
amniotes.
• Amniotes develop extra-embryonic
membranes to support the embryo.

98
 Amniote ExtraEmbryonic Membranes

• During amniote development, four

extraembryonic membranes form around
the embryo:
– The chorion outermost membrane / functions in gas
exchange.
– The amnion encloses the amniotic fluid.
– The yolk sac encloses the yolk.
– The allantois disposes of nitrogenous waste products
and contributes to gas exchange.

99
 Amnion
Allantois

Embryo
Amniotic Albumen
cavity
with
amniotic
fluid

Shell
Yolk
Chorion (nutrients)

Yolk sac 100

 Mammalian Development
• The eggs of placental mammals
– Are small yolk and store few nutrients
– Exhibit holoblastic cleavage
– Show no obvious polarity.
• Gastrulation and organogenesis resemble the
processes in birds and other reptiles.
• Early cleavage is relatively slow in humans
and other mammals.

101
• At completion of cleavage, the blastocyst
forms.
• A group of cells called the inner cell mass
develops into the embryo and forms the
extraembryonic membranes.
• The trophoblast, the outer epithelium of the
blastocyst, initiates implantation in the uterus,
and the inner cell mass of the blastocyst forms a
flat disk of cells.
• As implantation is completed, gastrulation
begins.

102
Early embryonic development of a human

Endometrial
epithelium
(uterine lining)

Uterus Inner cell mass

Trophoblast

Blastocoel

103
Early embryonic development of a human

Expanding
region of
trophoblast
Maternal
blood
vessel Epiblast

Hypoblast

Trophoblast

104
• The epiblast cells invaginate through a primitive streak 
to form mesoderm and endoderm.
• The placenta is formed from the trophoblast, 
mesodermal cells from the epiblast, and adjacent 
endometrial tissue.
• The placenta allows for the exchange of materials 
between the mother and embryo.
• By the end of gastrulation, the embryonic germ layers 
have formed.  The extraembryonic membranes in 
mammals are homologous to those of birds and other 
reptiles and develop in a similar way.

105
Early embryonic development of a
human Expanding
region of
trophoblast
Amniotic
cavity
Epiblast
Hypoblast
Yolk sac (from
hypoblast)
Extraembryonic
mesoderm cells
(from epiblast)
Chorion (from
trophoblast)
106
Early embryonic development of a human

Amnion

Chorion
Ectoderm
Mesoderm
Endoderm

Yolk sac

Extraembryonic
mesoderm

Atlantois

107
Four stages in early embryonic development of a human
Endometrial Expanding
epithelium region of
(uterine lining) trophoblast
Maternal
Uterus Inner cell mass blood
vessel Epiblast
Trophoblast
Hypoblast
Blastocoel Trophoblast

Expanding
region of
trophoblast Amnion
Amniotic
Chorion
cavity
Ectoderm
Epiblast
Mesoderm
Hypoblast
Endoderm
Yolk sac (from
hypoblast)
Yolk sac
Extraembryonic
mesoderm cells
(from epiblast) Extraembryonic
mesoderm
Chorion (from
trophoblast) Allantois

108
 Morphogenesis in animals involves
specific changes in cell shape, position,
and adhesion

• Morphogenesis is a major aspect of

development in plants and animals.
• Only in animals does it involve the
movement of cells.

109
 The Cytoskeleton, Cell Motility, and
Convergent Extension

• Changes in cell shape usually involve

reorganization of the cytoskeleton.
• Microtubules and microfilaments affect
formation of the neural tube.
• The cytoskeleton also drives cell migration,
or cell crawling, the active movement of
cells.
• In gastrulation, tissue invagination is
caused by changes in cell shape and
migration.
110
Ectoderm Change in cell
shape during
morphogenesis
Neural
plate

Microtubules

Actin filaments

Neural tube
111
 Role of Cell Adhesion
Molecules and the
Extracellular Matrix

• Cell adhesion molecules located on cell 
surfaces contribute to cell migration and stable 
tissue structure.
• One class of cell‐to‐cell adhesion molecule is 
the cadherins, which are important in 
formation of the frog blastula.

112
Cadherin is required for development of the blastula

RESULTS

0.25 mm 0.25 mm

Control embryo Embryo without EP cadherin

113
E-cadherin play a role in compaction

At compaction occurs, E-cadherin becomes restricted

to those sites on cell membranes that are in contact
with adjacent blastomeres.
114
 Compaction-
The first change of morphology

Uncompacted embryo Compacted embryo

(8-cell embryo) (Morula)
115
 The Mechanism of Compaction
Specific cell surface
proteins play a role in
compaction. One such
molecule, E-cadherin
(also known as
uvomorulin), a 120-kDa
adhesive glycoprotein, is
synthesized at the 2-cell
stage and is uniformly
spread throughout the
cell membrane.

In the first stage of compaction, each of the eight blastomeres

interacts with its neighbors to undergo membrane polarization. 116
 Formation of the
Vertebrate Limb
• The wings and legs of chicks, like all
vertebrate limbs, begin as bumps of tissue
called limb buds.
• The embryonic cells in a limb bud respond to
positional information indicating location
along three axes
– Proximal-distal axis
– Anterior-posterior axis
– Dorsal-ventral axis

117
 Anterior
Limb bud

Vertebrate limb AER

development Limb buds

Posterior
ZPA

50 µm

• Signal molecules produced

by inducing cells influence Apical
ectodermal
gene expression in cells ridge (AER)
receiving them.
• Signal molecules lead to (a) Organizer regions

differentiation and the 2

Digits
development of particular
structures. 3
4
Anterior
• Hox genes also play roles Ventral

during limb pattern formation. Proximal

Dorsal
Distal

Posterior

(b) Wing of chick embryo 118

 Knowledge Testing 1
1. Explain why most bryophytes grow close to
the ground and are restricted to periodically
moist environments.
2. Describe three traits that characterize modern
vascular plants and explain how these traits
have contributed to success on land

119
 Knowledge Testing 2
1. Explain why pollen grains were an important
adaptation for successful reproduction on
land
2. Diagram the generalized life cycle of an
angiosperm; indicate which structures are
part of the gametophyte generation and
which are part of the sporophyte generation.

120
 Knowledge Testing 3

1. List the characteristics that combine to define

animals
2. Compare the developmental differences
between protostomes and deuterostomes

121
 Knowledge Testing 4

1. Compare the formation of a blastula and

gastrulation in a sea urchin, a frog, and a chick
2. List and explain the functions of the
extraembryonic membranes
3. Explain pattern formation in a developing chick
limb.
4. Describe the role of the extracellular matrix in
embryonic development

122