Plant Diversity I How Plants Colonized Land: Powerpoint Lectures For

Chapter 29
Plant Diversity I
How Plants
Colonized Land
PowerPoint Lectures for
Biology, Seventh Edition
Neil Campbell and Jane Reece
Lectures by Chris Romero

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Overview: The Greening of Earth
• Looking at a lush landscape

– It is difficult to imagine the land without any
plants or other organisms
Figure 29.1
• For more than the first 3 billion years of Earth’s
history
– The terrestrial surface was lifeless
• Since colonizing land

– Plants have diversified into roughly 290,000
living species

• Concept 29.1: Land plants evolved from green
algae
• Researchers have identified green algae called
charophyceans as the closest relatives of land
plants

Morphological and Biochemical Evidence
• Many characteristics of land plants
– Also appear in a variety of algal clades

• There are four key traits that land plants share
only with charophyceans
– Rose-shaped complexes for cellulose
synthesis
Figure 29.2 30 nm
– Peroxisome enzymes
– Structure of flagellated sperm
– Formation of a phragmoplast

Genetic Evidence
• Comparisons of both nuclear and chloroplast
genes
– Point to charophyceans as the closest living
relatives of land plants
(a) Chara,
a pond 10 mm
organism
40 µm
(b) Coleochaete orbicularis, a disk-
Figure 29.3a, b shaped charophycean (LM)

Adaptations Enabling the Move to Land
• In charophyceans
– A layer of a durable polymer called
sporopollenin prevents exposed zygotes from
drying out
• The accumulation of traits that facilitated

survival on land
– May have opened the way to its colonization
by plants

• Concept 29.2: Land plants possess a set of
derived terrestrial adaptations
• Many adaptations
– Emerged after land plants diverged from their
charophycean relatives

Defining the Plant Kingdom
• Systematists
– Are currently debating the boundaries of the
plant kingdom
Viridiplantae
Streptophyta
Plantae
Red algae Chlorophytes Charophyceans Embryophytes
Figure 29.4 Ancestral alga

• Some biologists think that the plant kingdom
– Should be expanded to include some or all
green algae
• Until this debate is resolved

– This textbook retains the embryophyte
definition of kingdom Plantae

Derived Traits of Plants
• Five key traits appear in nearly all land plants
but are absent in the charophyceans
– Apical meristems
– Alternation of generations
– Walled spores produced in sporangia
– Multicellular gametangia
– Multicellular dependent embryos

• Apical meristems and alternation of
generations Apical
Developing
leaves Apical meristems of plant shoots
meristem
APICAL MERISTEMS of shoot
and roots. The light micrographs
are longitudinal sections at the tips
of a shoot and root.
Apical meristem
of root
Shoot Root
100 µm 100 µm
Haploid multicellular
organism (gametophyte)
Mitosis Mitosis
ALTERNATION OF GENERATIONS n
n n
Spores n n
Gametes
MEIOSIS FERTILIZATION
2n
2n Zygote
Mitosis
Diploid multicellular
Figure 29.5 organism (sporophyte)
Alternation of generations: a generalized scheme

• Walled spores; multicellular gametangia; and
multicellular, dependent embryos
Spores
WALLED SPORES PRODUCED IN SPORANGIA Sporangium
Sporophyte and sporangium Longitudinal section of

of Sphagnum (a moss) Sphagnum sporangium (LM)
Sporophyte
Gametophyte
MULTICELLULAR GAMETANGIA Female gametophyte

Archegonium
with egg
Antheridium
Archegonia and antheridia with sperm
of Marchantia (a liverwort)
Male
gametophyte
MULTICELLULAR, DEPENDENT EMBRYOS Embryo

Maternal tissue
2 µm
Embryo and placental

transfer cell of Marchantia
10 µm
Wall ingrowths
Figure 29.5 Placental transfer cell

• Additional derived units
– Such as a cuticle and secondary compounds,
evolved in many plant species

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
(a) Fossilized spores.
Unlike the spores of
most living plants,
which are single
grains, these spores
found in Oman are
in groups of four
(left; one hidden)
and two (right).
(b) Fossilized
sporophyte tissue.
The spores were
embedded in tissue
that appears to be
Figure 29.6 a, b from plants.

• Whatever the age of the first land plants
– Those ancestral species gave rise to a vast
diversity of modern plants
Table 29.1
• Land plants can be informally grouped
– Based on the presence or absence of vascular
tissue

• An overview of land plant evolution
Land plants
Vascular plants
Bryophytes
(nonvascular plants) Seedless vascular plants Seed plants
(ferns, horsetails, whisk fern)
Gymnosperms
(club mosses, spike mosses, quillworts)
Lycophytes
Pterophyte
Hornworts
Mosses
Angiosperms
Liverworts
Charophyceans
Origin of seed plants

(about 360 mya)
Origin of vascular
plants (about 420 mya)
Origin of land plants

(about 475 mya)
Ancestral
green alga
Figure 29.7
• Concept 29.3: The life cycles of mosses and
other bryophytes are dominated by the
gametophyte stage
• Bryophytes are represented today by three
phyla of small herbaceous (nonwoody) plants
– Liverworts, phylum Hepatophyta
– Hornworts, phylum Anthocerophyta
– Mosses, phylum Bryophyta

• Debate continues over the sequence of
bryophyte evolution
• Mosses are most closely related to vascular
plants

Bryophyte Gametophytes
• In all three bryophyte phyla
– Gametophytes are larger and longer-living
than sporophytes

• The life cycle of a moss
Raindrop
Key
Male
gametophyte Haploid (n)
1 Spores develop into
threadlike protonemata. Sperm Diploid (2n)
“Bud”
4 A sperm swims
Antheridia through a film of
2 The haploid moisture to an
protonemata 3 Most mosses have separate
archegonium and
produce “buds” male and female gametophytes,
fertilizes the egg.
that grow into with antheridia and archegonia,
gametophytes. respectively.
Protonemata “Bud”
Egg
Spores
Gametophore
Female Archegonia
8 Meiosis occurs and haploid
spores develop in the sporangium gametophyte
Peristome
of the sporophyte. When the
sporangium lid pops off, the Rhizoid
peristome “teeth” regulate 6 The sporophyte grows a
gradual release of the spores. long stalk, or seta, that emerges
Sporangium Seta from the archegonium. FERTILIZATION
MEIOSIS Capsule (within archegonium)
(sporangium) Calyptra Zygote
Mature
Mature
sporophytes
sporophytes Embryo
Foot
Archegonium
Young 5 The diploid zygote
sporophyte develops into a
Female sporophyte embryo within
Capsule with
peristome (LM)
gametophytes 7 Attached by its foot, the the archegonium.
sporophyte remains nutritionally
Figure 29.8 dependent on the gametophyte.

• Bryophyte gametophytes
– Produce flagellated sperm in antheridia
– Produce ova in archegonia
– Generally form ground-hugging carpets and

are at most only a few cells thick
• Some mosses
– Have conducting tissues in the center of their
“stems” and may grow vertically

Bryophyte Sporophytes
• Bryophyte sporophytes
– Grow out of archegonia
– Are the smallest and simplest of all extant

plant groups
– Consist of a foot, a seta, and a sporangium
• Hornwort and moss sporophytes

– Have stomata

• Bryophyte diversity
LIVERWORTS (PHYLUM HEPATOPHYTA)
Gametophore of
female gametophyte
Plagiochila
deltoidea,
Foot a “leafy”
Seta liverwort
Sporangium
Marchantia polymorpha,
a “thalloid” liverwort
500 µm
Marchantia sporophyte (LM)
HORNWORTS (PHYLUM ANTHOCEROPHYTA) MOSSES (PHYLUM BRYOPHYTA)

An Anthoceros Polytrichum commune,
hornwort species hairy-cap moss
Sporophyte
Sporophyte
Gametophyte
Gametophyte
Figure 29.9
Ecological and Economic Importance of Mosses
• Sphagnum, or “peat moss”
– Forms extensive deposits of partially decayed

organic material known as peat
– Plays an important role in the Earth’s carbon cycle
(a) Peat being harvested from a peat bog
Sporangium at
Gametophyte tip of sporophyte
Living
(b) Closeup of Sphagnum. Note the “leafy” gametophytes photo- Dead water-
and their offspring, the sporophytes. synthetic storing cells
cells 100 µm
(c) Sphagnum “leaf” (LM). The combination of living photosynthetic

cells and dead water-storing cells gives the moss its spongy quality.
(d)
“Tolland Man,” a bog mummy dating from 405–100 B.C.
The acidic, oxygen-poor conditions produced by
Sphagnum canpreserve human or other animal bodies for
Figure 29.10 a–d thousands of years.

• Concept 29.4: Ferns and other seedless
vascular plants formed the first forests
• Bryophytes and bryophyte-like plants
– Were the prevalent vegetation during the first
100 million years of plant evolution
• Vascular plants
– Began to evolve during the Carboniferous
period

Origins and Traits of Vascular Plants
• Fossils of the forerunners of vascular plants
– Date back about 420 million years

• These early tiny plants
– Had independent, branching sporophytes
– Lacked other derived traits of vascular plants
Figure 29.11
Life Cycles with Dominant Sporophytes
• In contrast with bryophytes
– Sporophytes of seedless vascular plants are
the larger generation, as in the familiar leafy
fern
– The gametophytes are tiny plants that grow on
or below the soil surface

• The life cycle of a fern
1 Sporangia release spores. 2 The fern spore 3 Although this illustration
Most fern species produce a single develops into a small, shows an egg and sperm
type of spore that gives rise to a photosynthetic gametophyte. from the same gametophyte,
Key bisexual gametophyte. a variety of mechanisms
Haploid (n) promote cross-fertilization
Diploid (2n) between gametophytes.
Antheridium
Spore Young
gametophyte
MEIOSIS
Sporangium
Archegonium Sperm
Mature Egg
New
sporophyte Zygote
sporophyte
Sporangium
FERTILIZATION
Sorus
6 On the underside
of the sporophyte‘s 4 Fern sperm use flagella
reproductive leaves to swim from the antheridia
Gametophyte
are spots called sori. to eggs in the archegonia.
Each sorus is a
cluster of sporangia.
Fiddlehead 5 A zygote develops into a new
sporophyte, and the young plant
grows out from an archegonium
Figure 29.12 of its parent, the gametophyte.

Transport in Xylem and Phloem
• Vascular plants have two types of vascular
tissue
– Xylem and phloem

• Xylem
– Conducts most of the water and minerals
– Includes dead cells called tracheids
• Phloem
– Distributes sugars, amino acids, and other
organic products
– Consists of living cells

Evolution of Roots
• Roots
– Are organs that anchor vascular plants
– Enable vascular plants to absorb water and

nutrients from the soil
– May have evolved from subterranean stems

Evolution of Leaves
• Leaves
– Are organs that increase the surface area of
vascular plants, thereby capturing more solar
energy for photosynthesis

• Leaves are categorized by two types
– Microphylls, leaves with a single vein
– Megaphylls, leaves with a highly branched

vascular system

• According to one model of evolution
– Microphylls evolved first, as outgrowths of
stems
Vascular tissue
(a) Microphylls, such as those of lycophytes, may have (b) Megaphylls, which have branched vascular
originated as small stem outgrowths supported by systems, may have evolved by the fusion of
Figure 29.13a, b single, unbranched strands of vascular tissue. branched stems.

Sporophylls and Spore Variations
• Sporophylls
– Are modified leaves with sporangia
• 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, having two types of spores
that give rise to male and female
gametophytes

Classification of Seedless Vascular Plants
• Seedless vascular plants form two phyla
– Lycophyta, including club mosses, spike
mosses, and quillworts
– Pterophyta, including ferns, horsetails, and
whisk ferns and their relatives

• The general groups of seedless vascular plants
LYCOPHYTES (PHYLUM LYCOPHYTA) Strobili

(clusters of
sporophylls)
Isoetes
gunnii,
a quillwort
Selaginella apoda,
a spike moss
Diphasiastrum tristachyum, a club moss
PTEROPHYTES (PHYLUM PTEROPHYTA)

Psilotum Equisetum Athyrium
nudum, arvense, filix-femina,
a whisk field lady fern
fern horsetail
Vegetative stem
Strobilus on
fertile stem
Figure 29.14 WHISK FERNS AND RELATIVES HORSETAILS FERNS

Phylum Lycophyta: Club Mosses, Spike Mosses, and
Quillworts
• Modern species of lycophytes

– Are relics from a far more eminent past
– Are small herbaceous plants

Phylum Pterophyta: Ferns, Horsetails, and Whisk
Ferns and Relatives
• Ferns
– Are the most diverse seedless vascular plants

The Significance of Seedless Vascular Plants
• The ancestors of modern lycophytes,
horsetails, and ferns
– Grew to great heights during the
Carboniferous, forming the first forests
Figure 29.15
• The growth of these early forests
– May have helped produce the major global
cooling that characterized the end of the
Carboniferous period
– Decayed and eventually became coal

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Chapter 29

Lectures by Chris Romero

• Looking at a lush landscape

• Since colonizing land

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Structure of flagellated sperm

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The accumulation of traits that facilitated

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Red algae Chlorophytes Charophyceans Embryophytes

Figure 29.4 Ancestral alga

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Until this debate is resolved

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Walled spores produced in sporangia

– Multicellular dependent embryos

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Sporophyte and sporangium Longitudinal section of

MULTICELLULAR GAMETANGIA Female gametophyte

MULTICELLULAR, DEPENDENT EMBRYOS Embryo

Embryo and placental

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

(ferns, horsetails, whisk fern)

Origin of seed plants

Origin of land plants

– Hornworts, phylum Anthocerophyta

– Mosses, phylum Bryophyta

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Produce ova in archegonia

– Generally form ground-hugging carpets and

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Are the smallest and simplest of all extant

• Hornwort and moss sporophytes

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

HORNWORTS (PHYLUM ANTHOCEROPHYTA) MOSSES (PHYLUM BRYOPHYTA)

– Forms extensive deposits of partially decayed

(c) Sphagnum “leaf” (LM). The combination of living photosynthetic

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Lacked other derived traits of vascular plants

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Includes dead cells called tracheids

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– Enable vascular plants to absorb water and

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Megaphylls, leaves with a highly branched

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings