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LECTURE PRESENTATIONS
For CAMPBELL BIOLOGY, NINTH EDITION
Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson
Chapter 29
Plant Diversity I: How Plants
Colonized Land
Lectures by
Erin Barley
Kathleen Fitzpatrick
© 2011 Pearson Education, Inc.
Overview: The Greening of Earth
• For more than the first 3 billion years of Earth’s
history, the terrestrial surface was lifeless
• Cyanobacteria likely existed on land 1.2 billion
years ago
• Around 500 million years ago, small plants,
fungi, and animals emerged on land
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• Since colonizing land, plants have diversified into
roughly 290,000 living species
• Land plants are defined as having terrestrial
ancestors, even though some are now aquatic
• Land plants do not include photosynthetic
protists (algae)
• Plants supply oxygen and are the ultimate
source of most food eaten by land animals
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Figure 29.1
1 m
Concept 29.1: Land plants evolved from
green algae
• Green algae called charophytes are the closest
relatives of land plants
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Morphological and Molecular Evidence
• Many characteristics of land plants also appear in
a variety of protist clades, mainly algae
• However, land plants share four key traits with
only charophytes
–
–
–
–
Rings of cellulose-synthesizing complexes
Peroxisome enzymes
Structure of flagellated sperm
Formation of a phragmoplast
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Figure 29.2
30 nm
1 m
• Comparisons of both nuclear and chloroplast
genes point to charophytes as the closest living
relatives of land plants
• Note that land plants are not descended from
modern charophytes, but share a common
ancestor with modern charophytes
© 2011 Pearson Education, Inc.
Figure 29.3
5 mm
Chara species, a pond organism
Coleochaete orbicularis, a
disk-shaped charophyte
that also lives in ponds (LM)
40 m
1 m
Figure 29.3a
5 mm
1 m
Chara species, a pond organism
Figure 29.3b
Coleochaete orbicularis, a
disk-shaped charophyte that
lives in ponds (LM)
40 m
1 m
Adaptations Enabling the Move to Land
• In charophytes a layer of a durable polymer called
sporopollenin prevents exposed zygotes from
drying out
• Sporopollenin is also found in plant spore walls
• The movement onto land by charophyte ancestors
provided 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
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• The accumulation of traits that facilitated survival
on land may have opened the way to its
colonization by plants
• Systematists are currently debating the
boundaries of the plant kingdom
• Some biologists think the plant kingdom should be
expanded to include some or all green algae
• Until this debate is resolved, we define plants as
embryophytes, plants with embryos
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Figure 29.4
Red algae
Chlorophytes
1 m
Plantae
Embryophytes
Streptophyta
Charophytes
Viridiplantae
ANCESTRAL
ALGA
Derived Traits of Plants
• Four key traits appear in nearly all land plants but
are absent in the charophytes
– Alternation of generations and multicellular,
dependent embryos
– Walled spores produced in sporangia
– Multicellular gametangia
– Apical meristems
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Alternation of Generations and Multicellular,
Dependent Embryos
• Plants alternate between two multicellular stages,
a reproductive cycle called alternation of
generations
• The 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
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• 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
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Figure 29.5a
Gametophyte
(n)
Mitosis
n
Gamete from
another plant
Mitosis
n
FERTILIZATION
2n
Sporophyte
(2n)
Haploid (n)
Diploid (2n)
n Spore Gamete n
MEIOSIS
Key
Zygote
Mitosis
Alternation of generations
1 m
Figure 29.5b
Embryo (LM) and
placental transfer cell (TEM)
of Marchantia (a liverwort)
Embryo
2 m
Maternal tissue
Wall ingrowths
10 m
1 m
Placental transfer
cell (outlined in
blue)
Figure 29.5ba
Embryo
Maternal tissue
10 m
1 m
Figure 29.5bb
2 m
Wall ingrowths
Placental transfer
cell (outlined in
blue)
1 m
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 makes
them resistant to harsh environments
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Figure 29.5c
Spores
Sporangium
Longitudinal section of
Sphagnum sporangium (LM)
Sporophyte
Gametophyte
1 m
Sporophytes and sporangia of Sphagnum (a moss)
Figure 29.5ca
Sporangium
Sporophyte
Gametophyte
Sporophytes and sporangia of Sphagnum (a moss)
1 m
Figure 29.5cb
Spores
Sporangium
Longitudinal section of
Sphagnum sporangium (LM)
1 m
Multicellular Gametangia
• Gametes are produced within organs called
gametangia
• Female gametangia, called archegonia, produce
eggs and are the site of fertilization
• Male gametangia, called antheridia, produce and
release sperm
© 2011 Pearson Education, Inc.
Figure 29.5d
Female
gametophyte
Archegonia,
each with an
egg (yellow)
Antheridia
(brown),
containing
sperm
Male
gametophyte
Archegonia and antheridia of Marchantia (a liverwort)
1 m
Figure 29.5da
Female
gametophyte
Male
gametophyte
1 m
Apical Meristems
• Plants sustain continual growth in their apical
meristems
• Cells from the apical meristems differentiate into
various tissues
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Figure 29.5e
Apical meristem
of shoot
Developing
leaves
Apical meristems of plant
roots and shoots
Apical
meristem
of root
Root
100 m
Shoot
1 m
100 m
Figure 29.5ea
Apical
meristem
of root
Root
100 m
1 m
Figure 29.5eb
Apical meristem
of shoot
Shoot
Developing
leaves
1 m
100 m
• Additional derived traits include
– Cuticle, a waxy covering of the epidermis
– Mycorrhizae, symbiotic associations between
fungi and land plants that may have helped plants
without true roots to obtain nutrients
– Secondary compounds that deter herbivores and
parasites
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The Origin and Diversification of Plants
• Fossil evidence indicates that plants were on land
at least 475 million years ago
• Fossilized spores and tissues have been extracted
from 475-million-year-old rocks
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Figure 29.6
(a) Fossilized
spores
(b) Fossilized
sporophyte
tissue
1 m
Figure 29.6a
(a) Fossilized
spores
Figure 29.6b
(b) Fossilized
sporophyte
tissue
• Those ancestral species gave rise to a vast
diversity of modern plants
© 2011 Pearson Education, Inc.
Figure 29.7
1 Origin of land plants (about 475 mya)
2 Origin of vascular plants (about 425 mya)
3 Origin of extant seed plants (about 305 mya)
Mosses
Land plants
ANCESTRAL
1
GREEN
ALGA
Nonvascular
plants
(bryophytes)
Liverworts
Hornworts
Pterophytes (ferns,
horsetails, whisk ferns)
3
Angiosperms
500
450
400
350
300
Millions of years ago (mya)
50
0
1 m
Seed plants
Gymnosperms
Vascular plants
2
Seedless
vascular
plants
Lycophytes (club
mosses, spike
mosses, quillworts)
Figure 29.7a
1 Origin of land plants (about 475 mya)
2 Origin of vascular plants (about 425 mya)
3 Origin of extant seed plants (about 305 mya)
Liverworts
ANCESTRAL
GREEN
1
ALGA
Mosses
Hornworts
Lycophytes (club
mosses, spike
mosses, quillworts)
2
Pterophytes (ferns,
horsetails, whisk ferns)
Gymnosperms
3
Angiosperms
500
450
350
400
300
Millions of years ago (mya)
50
1 m
0
Figure 29.7b
Land plants
Mosses
Nonvascular
plants
(bryophytes)
Liverworts
Hornworts
Angiosperms
Seed plants
Gymnosperms
1 m
Vascular plants
Pterophytes (ferns,
horsetails, whisk ferns)
Seedless
vascular
plants
Lycophytes (club
mosses, spike
mosses, quillworts)
• Land plants can be informally grouped based on
the presence or absence of vascular tissue
• Most plants have vascular tissue; these constitute
the vascular plants
• Nonvascular plants are commonly called
bryophytes
• Bryophytes are not a monophyletic group; their
relationships to each other and to vascular plants
are unresolved
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• Seedless vascular plants can be divided into
clades
– Lycophytes (club mosses and their relatives)
– Pterophytes (ferns and their relatives)
• Seedless vascular plants are paraphyletic, and are
of the same level of biological organization, or
grade
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• 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
– Angiosperms, the flowering plants
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Table 29. 1
1 m
Concept 29.2: Mosses and other nonvascular
plants have life cycles dominated by
gametophytes
• Bryophytes are represented today by three phyla
of small herbaceous (nonwoody) plants
– Liverworts, phylum Hepatophyta
– Hornworts, phylum Anthocerophyta
– Mosses, phylum Bryophyta
• Bryophyte refers to all nonvascular plants,
whereas Bryophyta refers only to the phylum of
mosses
© 2011 Pearson Education, Inc.
Figure 29.UN01
Nonvascular plants (bryophytes)
Seedless vascular plants
Gymnosperms
Angiosperms
1 m
Bryophyte Gametophytes
• In all three bryophyte phyla, gametophytes are
larger and longer-living than sporophytes
• Sporophytes are typically present only part of the
time
© 2011 Pearson Education, Inc.
Figure 29.8-1
“Bud”
Key
Haploid (n)
Diploid (2n)
Protonemata
(n)
“Bud”
Spores
Male
gametophyte
(n)
Gametophore
Spore
dispersal
Female
gametophyte
(n)
Peristome
Sporangium
MEIOSIS
Mature sporophytes
Rhizoid
Seta
Capsule
(sporangium)
2 mm
Foot
Capsule with
peristome (LM)
1 m
Female
gametophytes
Figure 29.8-2
“Bud”
Key
Haploid (n)
Diploid (2n)
Protonemata
(n)
“Bud”
Sperm
Antheridia
Male
gametophyte
(n)
Egg
Spores
Gametophore
Spore
dispersal
Female
gametophyte
(n)
Peristome
Sporangium
MEIOSIS
Mature sporophytes
Archegonia
Rhizoid
FERTILIZATION
(within archegonium)
Seta
Capsule
(sporangium)
2 mm
Foot
Capsule with
peristome (LM)
1 m
Female
gametophytes
Figure 29.8-3
“Bud”
Key
Haploid (n)
Diploid (2n)
Protonemata
(n)
“Bud”
Antheridia
Male
gametophyte
(n)
Sperm
Egg
Spores
Gametophore
Spore
dispersal
Female
gametophyte
(n)
Peristome
Sporangium
MEIOSIS
Mature sporophytes
Archegonia
Rhizoid
FERTILIZATION
Zygote (within archegonium)
(2n)
Seta
Capsule
(sporangium)
Foot
Embryo
2 mm
Archegonium
Capsule with
peristome (LM)
Young
sporophyte
(2n)
Female
gametophytes
1 m
2 mm
Figure 29.8a
1 m
Capsule with peristome (LM)
• A spore germinates into a gametophyte composed
of a protonema and gamete-producing
gametophore
• The height of gametophytes is constrained by lack
of vascular tissues
• Rhizoids anchor gametophytes to substrate
• Mature gametophytes produce flagellated sperm
in antheridia and an egg in each archegonium
• Sperm swim through a film of water to reach and
fertilize the egg
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Animation: Moss Life Cycle
© 2011 Pearson Education, Inc.
Bryophyte Sporophytes
• Bryophyte sporophytes grow out of archegonia,
and are the smallest and simplest sporophytes of
all extant plant groups
• A sporophyte consists of a foot, a seta (stalk),
and a sporangium, also called a capsule, which
discharges spores through a peristome
• Hornwort and moss sporophytes have stomata for
gas exchange; liverworts do not
© 2011 Pearson Education, Inc.
Figure 29.9a
Thallus
Gametophore of
female gametophyte
Sporophyte
Foot
Seta
Marchantia polymorpha,
a “thalloid” liverwort
Marchantia
sporophyte (LM)
500 m
Capsule
(sporangium)
Plagiochila deltoidea, a
“leafy” liverwort
1 m
Figure 29.9aa
Thallus
Gametophore of
female gametophyte
1 m
Marchantia polymorpha, a “thalloid” liverwort
Figure 29.9ab
Foot
Seta
Marchantia
sporophyte (LM)
500 m
Capsule
(sporangium)
1 m
Figure 29.9ac
Plagiochila
deltoidea,
a “leafy”
liverwort
1 m
Figure 29.9b
An Anthoceros
hornwort species
Sporophyte
Gametophyte
1 m
Figure 29.9c
Polytrichum commune,
hairy-cap moss
Capsule
Seta
Sporophyte
(a sturdy
plant that
takes months
to grow)
Gametophyte
1 m
The Ecological and Economic Importance
of Mosses
• Mosses are capable of inhabiting diverse and
sometimes extreme environments, but are
especially common in moist forests and wetlands
• Some mosses might help retain nitrogen in the
soil
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Figure 29.10
Annual nitrogen loss
(kg/ha)
RESULTS
6
5
4
3
2
1
0
With moss
Without moss
1 m
• Sphagnum, or “peat moss,” forms extensive
deposits of partially decayed organic material
known as peat
• Peat can be used as a source of fuel
• Sphagnum is an important global reservoir of
organic carbon
• Overharvesting of Sphagnum and/or a drop in
water level in peatlands could release stored CO2
to the atmosphere
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Figure 29.11
(a) Peat being harvested from a
peatland
(b) “Tollund Man,” a bog mummy
dating from 405–100 B.C.E.
1 m
Figure 29.11a
(a) Peat being harvested from1am
peatland
Figure 29.11b
(b) “Tollund Man,” a bog mummy dating from
405–100 B.C.E.
1 m
Concept 29.3: Ferns and other seedless
vascular plants were the first plants to
grow tall
• Bryophytes and bryophyte-like plants were the
prevalent vegetation during the first 100 million
years of plant evolution
• Vascular plants began to diversify during the
Devonian and Carboniferous periods
• Vascular tissue allowed these plants to grow tall
• Seedless vascular plants have flagellated sperm
and are usually restricted to moist environments
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Figure 29.UN03
Nonvascular plants (bryophytes)
Seedless vascular plants
Gymnosperms
Angiosperms
1 m
Origins and Traits of Vascular Plants
• Fossils of the forerunners of vascular plants date
back about 425 million years
• These early tiny plants had independent,
branching sporophytes
• Living vascular plants are characterized by
 Life cycles with dominant sporophytes
 Vascular tissues called xylem and phloem
 Well-developed roots and leaves
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Figure 29.12
Sporangia
1 m
Life Cycles with Dominant Sporophytes
• In contrast with bryophytes, sporophytes of
seedless vascular plants are the larger generation,
as in familiar ferns
• The gametophytes are tiny plants that grow on or
below the soil surface
Animation: Fern Life Cycle
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Figure 29.13-1
Key
Haploid (n)
Diploid (2n)
MEIOSIS
Spore
dispersal
Sporangium
Sporangium
Mature
sporophyte
(2n)
Sorus
Fiddlehead (young leaf)
1 m
Figure 29.13-2
Key
Haploid (n)
Diploid (2n)
MEIOSIS
Spore
dispersal
Spore
(n)
Rhizoid
Underside
of mature
gametophyte
(n)
Sporangium
Sporangium
Antheridium
Young
gametophyte
Archegonium
Egg
Mature
sporophyte
(2n)
FERTILIZATION
Sorus
Fiddlehead (young leaf)
Sperm
1 m
Figure 29.13-3
Key
Haploid (n)
Diploid (2n)
MEIOSIS
Spore
dispersal
Spore
(n)
Rhizoid
Underside
of mature
gametophyte
(n)
Sporangium
Sporangium
Antheridium
Young
gametophyte
Mature
sporophyte
(2n)
Sorus
New
sporophyte
Sperm
Archegonium
Egg
Zygote
(2n)
Gametophyte
Fiddlehead (young leaf)
1 m
FERTILIZATION
Transport in Xylem and Phloem
• Vascular plants have two types of vascular tissue:
xylem and phloem
• Xylem conducts most of the water and minerals
and includes dead cells called tracheids
• Water-conducting cells are strengthened by lignin
and provide structural support
• Phloem consists of living cells and distributes
sugars, amino acids, and other organic products
• Vascular tissue allowed for increased height,
which provided an evolutionary advantage
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Evolution of Roots
• Roots are organs that anchor vascular plants
• They enable vascular plants to absorb water and
nutrients from the soil
• Roots may have evolved from subterranean stems
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Evolution of Leaves
• Leaves are organs that increase the surface area
of vascular plants, thereby capturing more solar
energy that is used for photosynthesis
• Leaves are categorized by two types
 Microphylls, leaves with a single vein
 Megaphylls, leaves with a highly branched
vascular system
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• According to one model of evolution, microphylls
evolved as outgrowths of stems
• Megaphylls may have evolved as webbing
between flattened branches
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Figure 29.14
Overtopping
growth
Vascular tissue
Sporangia
Microphyll
Megaphyll
Other
stems
become
reduced
and
flattened.
(a) Microphylls
(b) Megaphylls
1 m
Webbing
develops.
Figure 29.14a
Vascular tissue
Sporangia
(a) Microphylls
1 m
Microphyll
Figure 29.14b
Overtopping
growth
Megaphyll
Other
stems
become
reduced
and
flattened.
(b) Megaphylls
Webbing
develops.
1 m
Sporophylls and Spore Variations
• Sporophylls are modified leaves with sporangia
• Sori are clusters of sporangia on the undersides
of sporophylls
• Strobili are cone-like structures formed from
groups of sporophylls
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• Most seedless vascular plants are homosporous,
producing one type of spore that develops into a
bisexual gametophyte
• All seed plants and some seedless vascular plants
are heterosporous
• Heterosporous species produce megaspores,
which give rise to female gametophytes, and
microspores, which give rise to male
gametophytes
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Classification of Seedless Vascular Plants
• There are two phyla of seedless vascular plants
– Phylum Lycophyta includes club mosses, spike
mosses, and quillworts
– Phylum Pterophyta includes ferns, horsetails, and
whisk ferns and their relatives
© 2011 Pearson Education, Inc.
Figure 29.15a
2.5 cm
1 cm
Selaginella
moellendorffii,
a spike moss
Isoetes
gunnii,
a quillwort
Strobili
(clusters of
sporophylls)
Diphasiastrum
1 m tristachyum,
a club moss
Selaginella
moellendorffii,
a spike moss
1 m
1 cm
Figure 29.15aa
Figure 29.15ab
Isoetes
gunnii,
a quillwort
1 m
Figure 29.15ac
2.5 cm
Strobili
(clusters of
sporophylls)
1 m
Diphasiastrum tristachyum,
a club moss
Figure 29.15b
Equisetum arvense,
field horsetail
Athyrium
filix-femina,
lady fern
1.5 cm
25 cm
Vegetative stem
4 cm
Psilotum
nudum,
a whisk
fern
Figure 29.15ba
1 m
25 cm
Athyrium
filix-femina,
lady fern
Figure 29.15bb
Equisetum arvense,
field horsetail
1.5 cm
Vegetative stem
Figure 29.15bc
4 cm
Psilotum
nudum,
a whisk
fern
1 m
Phylum Lycophyta: Club Mosses, Spike
Mosses, and Quillworts
• Giant lycophytes trees thrived for millions of years
in moist swamps
• Surviving species are small herbaceous plants
• Club mosses and spike mosses have vascular
tissues and are not true mosses
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Phylum Pterophyta: Ferns, Horsetails, and
Whisk Ferns and Relatives
• Ferns are the most diverse seedless vascular
plants, with more than 12,000 species
• They are most diverse in the tropics but also thrive
in temperate forests
• Horsetails were diverse during the Carboniferous
period, but are now restricted to the genus
Equisetum
• Whisk ferns resemble ancestral vascular plants
but are closely related to modern ferns
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The Significance of Seedless Vascular Plants
• The ancestors of modern lycophytes, horsetails,
and ferns grew to great heights during the
Devonian and Carboniferous, forming the first
forests
• Increased growth and photosynthesis removed
CO2 from the atmosphere and may have
contributed to global cooling at the end of the
Carboniferous period
• The decaying plants of these Carboniferous
forests eventually became coal
© 2011 Pearson Education, Inc.
Figure 29.16
Fern
Lycophyte trees
Horsetail
Tree trunk
covered with
small leaves
1 m
Lycophyte tree
reproductive
structures
Figure 29.UN02
1 m
Figure 29.UN04
Homosporous spore production
Sporangium
on sporophyll
Single
type of spore
Typically a
bisexual
gametophyte
Eggs
Sperm
Heterosporous spore production
Megasporangium
on megasporophyll
Megaspore
Female
gametophyte
Eggs
Microsporangium
on microsporophyll
Microspore
Male
gametophyte
Sperm
1 m
Figure 29.UN05
Gametophyte
Mitosis
Mitosis
n
n
n
Spore
n
Gamete
MEIOSIS
Apical meristem
of shoot
Developing
leaves
FERTILIZATION
2n Zygote
Haploid
Diploid
Mitosis
Sporophyte
1 Alternation of generations
Archegonium
with egg
2 Apical meristems
Antheridium
with sperm
Sporangium
Spores
1 m
3 Multicellular gametangia
4 Walled spores in sporangia
Figure 29.UN05a
Gametophyte
Mitosis
Mitosis
n
n
n
Spore
n
Gamete
MEIOSIS
FERTILIZATION
2n Zygote
Haploid
Diploid
Mitosis
Sporophyte
1 Alternation of generations
1 m
Figure 29.UN05b
Apical meristem
of shoot
Developing
leaves
2 Apical meristems
1 m
Figure 29.UN05c
Archegonium
with egg
Antheridium
with sperm
3 Multicellular gametangia
1 m
Figure 29.UN05d
Sporangium
Spores
4 Walled spores in sporangia
1 m
Figure 29.UN06
1 m
Figure 29.UN07
1 m
Figure 29.UN08
1 m
Figure 29.UN09
1 m
Figure 29.UN10
1 m