Chapter 29 Plant Diversity I: How Plants Colonized Land

Lecture Outline

Overview: The Greening of Earth

 For the first 3 billion years of Earth’s history, the land was
lifeless.
 Thin coatings of cyanobacteria existed on land about 1.2
billion years ago.
 About 500 million years ago, plants, fungi, and animals joined
them.
 More than 290,000 species of plants inhabit Earth today.
 Most plants live in terrestrial environments, including
deserts, grasslands, and forests.
 Some species, such as sea grasses, have returned to aquatic
habitats.
 The presence of plants has enabled other organisms to
survive on land.
 Plant roots have created habitats for other organisms by
stabilizing landscapes.
 Plants are the source of oxygen and the ultimate provider
of food for land animals.

Concept 29.1 Land plants evolved from green algae

 Researchers have identified a lineage of green algae called
charophyceans as the closest relatives of land plants.
 Many key characteristics of land plants also appear in a
variety of algal clades.
 Plants are multicellular, eukaryotic, photosynthetic
autotrophs.
 But red, brown, and some green algae also fit this
description.
 Plants have cell walls made of cellulose.
 So do green algae, dinoflagellates, and brown algae.
 Plants have chloroplasts with chlorophyll a and b.
 So do green algae, euglenids, and a few dinoflagellates.
 Land plants share four key features only with the
charophyceans.
1. The plasma membranes of land plants and
charophyceans possess rosette cellulose-synthesizing
complexes that synthesize the cellulose microfibrils of the
cell wall.
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  These complexes contrast with the linear arrays of
cellulose-producing proteins in noncharophycean algae.
 Also, the cell walls of plants and charophyceans contain a
higher percentage of cellulose than the cell walls of
noncharophycean algae.
2. A second feature that unites charophyceans and land
plants is the presence of peroxisome enzymes to help
minimize the loss of organic products as a result of
photorespiration.
 Peroxisomes of other algae lack these enzymes.
3. In those land plants that have flagellated sperm cells,
the structure of the sperm resembles the sperm of
charophyceans.
4. Finally, certain details of cell division are common only
to land plants and the most complex charophycean algae.
 These include the formation of a phragmoplast, an alignment
of cytoskeletal elements and Golgi-derived vesicles, during
the synthesis of new cross-walls during cytokinesis.
 Over the past decade, researchers involved in an
international initiative called “Deep Green” have conducted a
large-scale study of the major transitions in plant evolution.
 These researchers have analyzed genes from a wide
range of plant and algal species.
 Comparisons of nuclear and chloroplast genes support the
hypothesis that the charophyceans are the closest living
relatives of land plants.
 Many charophycean algae inhabit shallow waters at the
edges of ponds and lakes, where they experience occasional
drying.
 In such environments, natural selection favors individuals
that can survive periods when they are not submerged in
water.
 A layer of a durable polymer called sporopollenin prevents
exposed charophycean zygotes from drying out until they
are in water again.
 This chemical adaptation may have been the precursor to
the tough sporopollenin walls that encase plant spores.
 The accumulation of such traits by at least one population
of ancestral charophyceans enabled their descendents—the
first land plants—to live permanently above the waterline.
 The evolutionary novelties of the first land plants opened an
expanse of terrestrial habitat previously occupied only by
films of bacteria.
 The new frontier was spacious.
 The bright sunlight was unfiltered by water and plankton.
 The atmosphere had an abundance of carbon dioxide.
 The soil was rich in mineral nutrients.
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  At least at first, there were relatively few herbivores or
pathogens.

Concept 29.2 Land plants possess a set of derived terrestrial

adaptations
 A number of adaptations evolved in plants that allowed them
to survive and reproduce on land.
 What exactly is the line that divides land plants from algae?
 We will adopt the traditional scheme, which equates the
kingdom Plantae with embryophytes (plants with embryos).
 Some botanists now propose that the plant kingdom should
be renamed the kingdom Streptophyta and expanded to
include the charophyceans and a few related groups.
 Others suggest the kingdom Viridiplantae, which includes
chlorophytes as well as plants.
 Five key traits appear in nearly all land plants but are
absent in the charophyceans.
 We infer that these traits evolved as derived traits of land
plants.
 The five traits are:
1. Apical meristems.
2. Alternation of generations.
3. Multicellular embryo that is dependent on the parent
plant.
4. Sporangia that produce walled spores.
5. Gametangia that produce gametes.
Apical meristems
 In terrestrial habitats, the resources that a photosynthetic
organism requires are found in two different places.
 Light and carbon dioxide are mainly aboveground.
 Water and mineral resources are found mainly in the soil.
 Therefore, plants show varying degrees of structural
specialization for subterranean and aerial organs—roots and
shoots in most plants.
 The elongation and branching of the shoots and roots
maximize their exposure to environmental resources.
 This growth is sustained by apical meristems, localized
regions of cell division at the tips of shoots and roots.
 Cells produced by meristems differentiate into various
tissues, including surface epidermis and internal tissues.
Alternation of generations
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  All land plants show alternation of generations in which two
multicellular body forms alternate.
 This life cycle also occurs in various algae.
 However, alternation of generations does not occur in the
charophyceans, the algae most closely related to land plants.
 In alternation of generations, one of the multicellular
bodies is called the gametophyte and has haploid cells.
 Gametophytes produce gametes, egg and sperm, by mitosis.
 Fusion of egg and sperm during fertilization form a diploid
zygote.
 Mitotic division of the diploid zygote produces the other
multicellular body, the sporophyte.
 Meiosis in a mature sporophyte produces haploid
reproductive cells called spores.
 A spore is a reproductive cell that can develop into a new
organism without fusing with another cell.
 Mitotic division of a plant spore produces a new multicellular
gametophyte.
 Unlike the life cycles of other sexually producing organisms,
alternation of generations in land plants (and some algae)
results in both haploid and diploid stages that exist as
multicellular bodies.
 For example, humans do not have alternation of generations
because the only haploid stage in the life cycle is the
gamete, which is single-celled.
Walled spores produced by sporangia
 Plant spores are haploid reproductive cells that grow into
gametophytes by mitosis.
 Sporopollenin makes the walls of spores very tough and
resistant to harsh environments.
 Multicellular organs called sporangia are found on the
sporophyte and produce spores.
 Within sporangia, diploid cells called sporocytes undergo
meiosis and generate haploid spores.
 The outer tissues of the sporangium protect the developing
spores until they are ready to be released into the air.
Multicellular gametangia
 Plant gametophytes produce gametes within multicellular
organs called gametangia.
 A female gametangium, called an archegonium, produces a
single egg cell in a vase-shaped organ.
 The egg is retained within the base.
 Male gametangia, called antheridia, produce and release
sperm into the environment.
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  In many major groups of living plants, the sperm have
flagella and swim to the eggs though a water film.
 Each egg is fertilized within an archegonium, where the
zygote develops into the embryo.
 The gametophytes of seed plants are so reduced in size
that archegonia and antheridia have been lost in some lineages.
Multicellular, dependent embryos
 Multicellular plant embryos develop from zygotes that are
retained within tissues of the female parent.
 The multicellular, dependent embryo of land plants is such a
significant derived trait that land plants are also known as
embryophytes.
 The parent provides nutrients, such as sugars and amino
acids, to the embryo.
 The embryo has specialized placental transfer cells that
enhance the transfer of nutrients from parent to embryo.
 These are sometimes present in the adjacent maternal
tissues as well.
 This interface is analogous to the nutrient-transferring
embryo-mother interface of placental mammals.
 Additional derived traits have evolved in many plant species.
 The epidermis of many plants has a cuticle consisting of
polymers called polyesters and waxes.
 The cuticle waterproofs the epidermis, preventing excessive
water loss, and offers protection from microbial attack.
 Many land plants produce secondary compounds, so named
because they are the products of secondary metabolic
pathways that branch from primary metabolic pathways.
 Alkaloids, terpenes, and tannins defend against herbivores
and parasites.
 Flavonoids absorb harmful UV radiation and may act as
signals in symbiotic relationships with beneficial soil
microbes.
 Phenolics deter attack by pathogenic microbes.
Land plants have diversified since their origin from algal
ancestors.
 Fossils of plant spores have been extracted from 475-
million-year-old rocks in Oman.
 These spores were embedded in plant cuticle material that
is similar to spore-bearing tissue in living plants.
 These fossils clearly belong to plants.
 A 2001 study of the “molecular clock” of plants suggests
that the common ancestor of living plants existed 700 million
years ago.
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  A 2003 study suggests a new date of 490 to 425 million
years, roughly the same age as the spores found in Oman.
 Land plants can be informally grouped based on the
presence or absence of an extensive system of vascular
tissue, cells joined into tubes that transport water and
nutrients throughout the plant body.
 Plants that do not have an extensive transport system are
described as “nonvascular plants,” although some mosses do
have simple vascular tissue.
 Nonvascular plants are informally called bryophytes.
 There is some uncertainty about whether or not bryophytes
are monophyletic and represent a clade.
 Vascular plants form a clade consisting of 93% of all land
plants.
 Three smaller clades are found within the vascular plants.
 Lycophytes include club mosses and their relatives.
 Pterophytes include the ferns and their relatives.
 These two clades are called the seedless vascular
plants.
 A third clade of vascular plants includes the seed plants,
the vast majority of living plants.
 A seed is an embryo packaged with a supply of nutrients
within a protective coat.
 Seed plants can be divided into two groups: gymnosperms
and angiosperms.
 Gymnosperms are called “naked seed plants” because their
seeds are not enclosed in chambers.
 Angiosperms are a huge clade including all flowering plants.

Concept 29.3 The life cycles of mosses and other bryophytes

are dominated by the gametophyte stage
 Bryophytes are represented by three phyla:
 Phylum Hepatophyta—liverworts
 Phylum Anthocerophyta—hornworts
 Phylum Bryophyta—mosses
 Note that the name Bryophyta refers only to one phylum,
but the informal term bryophyte refers to all nonvascular
plants.
 It has not been established whether the diverse bryophytes
form a clade.
 Systematists continue to debate the sequence in which the
three phyla of bryophytes evolved.

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  Bryophytes acquired many unique adaptations after their
evolutionary split from the ancestors of modern vascular
plants.
 They also possess some ancestral traits characteristic of
the earliest plants.
 In bryophytes, gametophytes are the largest and most
conspicuous phase of the life cycle.
 Sporophytes are smaller and are present only part of the
time.
 Bryophyte spores germinate in favorable habitats and grow
into gametophytes by mitosis.
 The gametophyte is a mass of green, branched, filaments
that are one cell thick, called a protonema.
 A protonema has a large surface area that enhances
absorption of water and minerals.
 In favorable conditions, protonema generate gamete-
producing structures, the gametophores.
 Bryophytes are anchored by tubular cells or filaments of
cells, called rhizoids.
 Unlike roots, rhizoids are not composed of tissues, lack
specialized conducting cells, and do not play a primary role in
water and mineral absorption.
 Bryophyte gametophytes are generally only one or a few
cells thick, placing all cells close to water and dissolved
minerals.
 Most bryophytes lack conducting tissues to distribute water
and organic compounds within the gametophyte.
 Some mosses have conducting tissues in their stems, and a
few can grow as tall as 2 m.
 It is not clear if conducting tissues in mosses are analogous
or homologous to the xylem and phloem of vascular plants.
 Lacking support tissues, most bryophytes are only a few
centimeters tall.
 The mature gametophores of bryophytes produce gametes
in gametangia.
 Each vase-shaped archegonium produces a single egg.
 Elongated antheridia produce many flagellated sperm.
 When plants are coated with a thin film of water, sperm
swim toward the archegonia, drawn by chemical attractants.
 They swim into the archegonia and fertilize the eggs.
 The zygotes and young sporophytes are retained and
nourished by the parent gametophyte.
 Layers of placental nutritive cells transport materials from
parent to embryos.

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 Bryophyte sporophytes disperse enormous numbers of spores.
 While the bryophyte sporophyte does have photosynthetic
plastids when young, it cannot live apart from the maternal
gametophyte.
 A bryophyte sporophyte remains attached to its maternal
gametophyte throughout the sporophyte’s lifetime.
 It depends on the gametophyte for sugars, amino acids,
minerals, and water.
 Bryophytes have the smallest and simplest sporophytes of
all modern plant groups, consistent with the hypothesis that
larger and more complex sporophytes evolved only later in
vascular plants.
 Moss sporophytes consist of a foot, an elongated stalk (the
seta), and a sporangium (the capsule).
 The foot gathers nutrients and water from the parent
gametophyte via transfer cells.
 The stalk conducts these materials to the capsule.
 In most mosses, the seta becomes elongated, elevating the
capsule and enhancing spore dispersal.
 The moss capsule (sporangium) is the site of meiosis and
spore production.
 One capsule can generate more than 50 million spores.
 When immature, the capsule is covered by a protective cap
of gametophyte tissue, the calyptra.
 This is lost when the capsule is ready to release spores.
 The upper part of the capsule, the peristome, is often
specialized for gradual spore release.
 Liverworts have the simplest sporophytes among the
bryophytes.
 They consist of a short stalk bearing round sporangia that
contain the developing spores, and a nutritive foot
embedded in gametophyte tissues.
 Hornwort and moss sporophytes are larger and more
complex.
 Hornwort sporophytes resemble grass blades and have a
cuticle.
 The sporophytes of mosses start out green and
photosynthetic, but turn tan or brownish red when ready to
release their spores.
 The sporophytes of hornworts and mosses have epidermal
stomata, like those of vascular plants.
 These pores support photosynthesis by allowing the
exchange of CO2 and O2 between the outside air and the
interior of the sporophyte.
 The fact that stomata are present in mosses and hornworts
but absent in liverworts has led to three hypotheses for
their evolution.
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  If liverworts are the deepest-branching lineage of
land plants, then stomata evolved once in the ancestor of
hornworts, mosses and vascular plants.
 If hornworts are the deepest-branching lineage of
land plants, then stomata evolved once and were lost in
the liverwort lineage.
 Perhaps hornworts acquired stomata independently of
mosses and vascular plants.
Bryophytes provide many ecological and economic benefits.
 Wind dispersal of lightweight spores has distributed
bryophytes around the world.
 They are common and diverse in moist forests and wetlands.
 Some even inhabit extreme environments such as
mountaintops, tundra, and deserts.
 Phenolic compounds in moss cell walls absorb damaging levels
of radiation present in deserts and at high altitudes and
latitudes.
 Many mosses can exist in very cold or dry habitats because
they are able to lose most of their body water and then
rehydrate and reactivate their cells when moisture again
becomes available.
 Few vascular plants can survive the same degree of
desiccation.
 Sphagnum, a wetland moss, is especially abundant and
widespread.
 It forms extensive deposits of undecayed organic material,
called peat.
 Wet regions dominated by Sphagnum or peat moss are
known as peat bogs.
 Its organic materials do not decay readily because of
resistant phenolic compounds and acidic secretions that
inhibit bacterial activity.
 Peatlands, extensive high-latitude boreal wetlands occupied
by Sphagnum, play an important role as carbon reservoirs,
stabilizing atmospheric carbon dioxide levels.
 Sphagnum has been used in the past for diapers and as a
natural antiseptic material for wounds.
 Today, it is harvested for use as a soil conditioner and for
packing plants’ roots because of the water storage capacity of
its large, dead cells.
 Worldwide, an estimated 400 billion tons of organic carbon
are stored as peat.

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Concept 29.4 Ferns and other seedless vascular plants formed
the first forests
Ferns and other seedless vascular plants flourished in the
Carboniferous period.
 Bryophytes were the prevalent vegetation for the first 100
million years that terrestrial communities existed.
 Then vegetation began to take on a taller profile with the
evolution of vascular plants.
 Modern seedless vascular plants provide insights into plant
evolution during the Carboniferous period, when vascular plants
began to diversify, but most groups of seed plants had not yet
evolved.
 The sperm of ferns and all other seedless vascular plants
are flagellated and must swim through a film of water to reach
eggs.
 Due to the swimming sperm and their fragile gametophytes,
modern seedless vascular plants are most common in damp
environments.
 Fossils of the ancestors of today’s vascular plants date
back about 420 million years.
 Unlike bryophytes, these plants had branched sporophytes
that did not remain dependent on gametophytes for growth.
Five main traits characterize modern vascular plants.
 Five main traits characterize modern vascular plants:
1. Life cycles with dominant sporophytes.
2. Transport in xylem and phloem.
3. Evolution of roots.
4. Evolution of leaves.
5. Sporophylls and spore variations.
Life cycles with dominant sporophytes
 Fossils suggest that the ancestors of vascular plants had
life cycles characterized by gametophytes and sporophytes
that were about equal in size.
 Among living vascular plants, the sporophyte generation is
the larger and more complex plant.
 For example, the leafy fern plants that you are familiar with
are sporophytes.
 The gametophytes are tiny plants that grow on or just below
the soil surface.

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  This reduction in the size of the gametophytes is even more
extreme in seed plants.
Transport in xylem and phloem
 Vascular plants have two types of vascular tissue: xylem and
phloem.
 Xylem conducts most of the water and minerals.
 The xylem of all vascular plants includes tracheids, tube-
shaped cells that carry water and minerals up from roots.
 When functioning, these cells are dead, with only their walls
providing a system of microscopic water pipes.
 The water-conducting cells in vascular plants are lignified,
strengthened by the phenolic polymer lignin.
 Phloem is a living tissue in which nutrient-conducting cells
are arranged into tubes that distribute sugars, amino acids,
and other organic products.
 Lignified vascular tissue permitted vascular plants to grow
to greater heights than bryophytes.
Evolution of roots
 Lignified vascular tissue also allowed the evolution of roots.
 Roots are organs that anchor vascular plants and enable
them to absorb water and nutrients from the soil.
 Roots also allow the shoot system to grow taller.
 Roots may have evolved from the subterranean portions of
stems in ancient vascular plants.
 It is not clear whether roots evolved once in the common
ancestor of all vascular plants or independently in different
lineages.
 Studying genes that control root development may resolve
this controversy.
Evolution of leaves
 Leaves are organs that increase the surface area of
vascular plants, capturing more solar energy for
photosynthesis.
 In terms of size and complexity, leaves can be classified as
microphylls and megaphylls.
 All lycophytes have microphylls, small leaves with only a
single unbranched vein.
 These leaves probably evolved as small outgrowths on the
surface of stems, supported by single strands of vascular
tissue.
 All other vascular plants have megaphylls, leaves with a
highly branched vascular system.
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  A branched vascular system can deliver water and minerals
to the expanded leaf.
 It can also export larger quantities of sugars from the leaf.
 Megaphylls support more photosynthetic activity.
 The fossil evidence suggests that megaphylls evolved from a
series of branches lying close together on a stem.
 One hypothesis proposes that megaphylls evolved when the
branch system flattened and a tissue webbing developed,
joining the branches.
 Under this hypothesis, true, branched stems preceded the
origin of large leaves and roots.
Sporophylls and spore variations
 Vascular plants have sporophylls, modified plants that bear
sporangia.
 Sporophylls vary greatly in structure.
 Ferns produce clusters of sporangia called sori, usually on
the underside of leaves.
 In gymnosperms, groups of sporophylls form cone or
strobili.
 Another key variation among vascular plants is the
distinction between homosporous and heterosporous species.
 Most seedless vascular plants are homosporous, producing a
single type of spore.
 This spore develops into a bisexual gametophyte with both
archegonia (female sex organs) and antheridia (male sex
organs).
 Most ferns are homosporous.
 A heterosporous species produces two kinds of spores.
 Megaspores develop into female gametophytes.
 Microspores develop into male gametophytes.
 All seed plants and a few seedless vascular plants are
heterosporous.
Classification of seedless vascular plants.
 Living seedless vascular plants form two clades: lycophytes
and pterophytes.
 Lycophytes include club mosses, spike mosses, and
quillworts.
 Pterophytes include ferns, horsetails, whisk ferns, and their
relatives.
 Phylum Lycophyta: club mosses, spike mosses, and
quillworts
 Modern species of lycophytes are relicts of an eminent past.

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  By the Carboniferous, there were two evolutionary lineages
of lycophytes: small, herbaceous plants and giant, woody
trees standing 40 meters tall.
 The giant lycophytes became extinct as the climate cooled
and dried at the end of the Carboniferous.
 The small lycophytes survived, and are now represented by
1,200 species.
 Phylum Pterophyta: ferns, horsetails, and whisk ferns
 Ferns radiated extensively from their Devonian origins and
grew with lycophytes and horsetails in the Carboniferous
swamp forests.
 There are 12,000 species of living ferns.
 They are most diverse in the tropics, thrive in temperate
forests, and some can even survive arid conditions.
 Horsetails grew up to 15 meters in height during the
Carboniferous period.
 Today, only 15 species of single genus Equisetum survives.
 Psilotum, the whisk fern, and a close relative form a clade
of terrestrial epiphytes.
 Whisk ferns are the only vascular plants lacking true
roots and leaves.
 These plants have been considered “living fossils”
because their dichotomous branching and lack of true
leaves and roots seemed similar to early vascular plants.
 However, comparisons of DNA sequences and details of
sperm ultrastructure indicate that they are closely related
to ferns.
 The whisk fern’s ancestors lost true leaves and roots during
evolution.
The significance of seedless vascular plants.
 The ancestors of modern lycophytes and ferns, along with
their seedless vascular relatives, formed the first forests
during the Carboniferous.
 With the evolution of vascular tissue, roots, and leaves,
these plants accelerated their rate of photosynthesis and
dramatically increased the removal of CO2 in the atmosphere.
 Scientists estimate that CO2 levels dropped by as much as a
factor of five during the Carboniferous, causing global cooling
and widespread glacier formation.
 The first forests gave rise to modern-day coal.
 In the stagnant waters of the Carboniferous, dead plants
did not fully decay.
 The organic material turned to thick layers of peat. Marine
sediments piled up on top, and over millions of years, heat and
pressure converted the peat to coal.
 Humans still burn 6 billion tons of coal each year.
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