plant diversity ii: the evolution of seed...
TRANSCRIPT
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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
© 2011 Pearson Education, Inc.
Lectures by
Erin Barley
Kathleen Fitzpatrick
Plant Diversity II: The Evolution
of Seed Plants
Chapter 30 Overview: Transforming the World
• Seeds changed the course of plant evolution,
enabling their bearers to become the dominant
producers in most terrestrial ecosystems
• Seed plants originated about 360 million years
ago
• A seed consists of an embryo and nutrients
surrounded by a protective coat
• Domestication of seed plants had begun by
8,000 years ago and allowed for permanent
settlements
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Figure 30.1
Concept 30.1: 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
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Advantages of Reduced Gametophytes
• The gametophytes of seed plants develop within
the walls of spores that are retained within
tissues of the parent sporophyte
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Figure 30.2
PLANT GROUP
Mosses and other
nonvascular plants
Gametophyte
Sporophyte
Sporophyte
(2n)
Gametophyte
(n)
Dominant
Reduced, dependent on
gametophyte for nutrition Dominant
Reduced, independent
(photosynthetic and
free-living)
Ferns and other seedless
vascular plants
Dominant
Reduced (usually microscopic), dependent on surrounding
sporophyte tissue for nutrition
Seed plants (gymnosperms and angiosperms)
Angiosperm Gymnosperm
Microscopic
female
gametophytes
(n) inside
these parts
of flowers
Microscopic female
gametophytes (n) inside
ovulate cone
Microscopic
male
gametophytes
(n) inside
these parts
of flowers
Microscopic male
gametophytes (n)
inside pollen
cone
Sporophyte (2n) Sporophyte (2n)
Sporophyte
(2n)
Gametophyte
(n)
Example
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Figure 30.2a
Mosses and other
nonvascular plants
Gametophyte
Sporophyte
Sporophyte
(2n)
Gametophyte
(n)
Dominant
Reduced, dependent on
gametophyte for nutrition
Example
Figure 30.2b
Dominant
Reduced, independent
(photosynthetic and
free-living)
Ferns and other seedless vascular plants
Sporophyte
(2n)
Gametophyte
(n)
Gametophyte
Sporophyte
Example
Dominant
Reduced (usually microscopic), dependent on surrounding
sporophyte tissue for nutrition
Seed plants (gymnosperms and angiosperms)
Angiosperm Gymnosperm
Microscopic
female
gametophytes
(n) inside
these parts
of flowers
Microscopic female
gametophytes (n) inside
ovulate cone
Microscopic
male
gametophytes
(n) inside
these parts
of flowers
Microscopic male
gametophytes (n)
inside pollen
cone Sporophyte
(2n) Sporophyte
(2n)
Gametophyte
Sporophyte
Example
Figure 30.2c
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
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Ovules and Production of Eggs
• An ovule consists of a megasporangium,
megaspore, and one or more protective
integuments
• Gymnosperm megaspores have one integument
• Angiosperm megaspores usually have two
integuments
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Figure 30.3-1
Immature
ovulate cone
Integument (2n)
Spore wall
Megaspore (n)
(a) Unfertilized ovule
Megasporangium
(2n)
Pollen grain (n) Micropyle
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Pollen and Production of Sperm
• Microspores develop into pollen grains, which contain the male gametophytes
• Pollination is the transfer of pollen to the part of a seed plant containing the ovules
• Pollen eliminates the need for a film of water and can be dispersed great distances by air or animals
• If a pollen grain germinates, it gives rise to a pollen tube that discharges sperm into the female gametophyte within the ovule
© 2011 Pearson Education, Inc.
Figure 30.3-2
Immature
ovulate cone
Integument (2n)
Spore wall
Megaspore (n)
Female
gametophyte (n)
Egg nucleus
(n)
Discharged sperm nucleus (n)
Pollen tube
Male gametophyte (n)
(a) Unfertilized ovule
Megasporangium
(2n)
Pollen grain (n) Micropyle
(b) Fertilized ovule
The Evolutionary Advantage of Seeds
• A seed develops from the whole ovule
• A seed is a sporophyte embryo, along with its
food supply, packaged in a protective coat
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Figure 30.3-3
Immature
ovulate cone
Integument (2n)
Spore wall
Megaspore (n)
Female
gametophyte (n)
Egg nucleus
(n)
Discharged sperm nucleus (n)
Pollen tube
Male gametophyte (n)
(a) Unfertilized ovule
Megasporangium
(2n)
Pollen grain (n) Micropyle
(b) Fertilized ovule (c) Gymnosperm seed
Seed coat
Spore wall
Food supply (n)
Embryo (2n)
• Seeds provide some evolutionary advantages
over spores
– They may remain dormant for days to years,
until conditions are favorable for germination
– Seeds have a supply of stored food
– They may be transported long distances by
wind or animals
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Concept 30.2: Gymnosperms bear
“naked” seeds, typically on cones
• Gymnosperms means “naked seeds”
• The seeds are exposed on sporophylls that form
cones
• Angiosperm seeds are found in fruits, which are
mature ovaries
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Figure 30.UN01
Nonvascular plants (bryophytes)
Seedless vascular plants
Gymnosperms
Angiosperms
Gymnosperm Evolution
• Fossil evidence reveals that by the late Devonian
period some plants, called progymnosperms,
had begun to acquire some adaptations that
characterize seed plants
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Figure 30.4
• Living seed plants can be divided into two clades:
gymnosperms and angiosperms
• Gymnosperms appear early in the fossil record
about 305 million years ago and dominated
Mesozoic (251–65 million years ago) terrestrial
ecosystems
• Gymnosperms were better suited than
nonvascular plants to drier conditions
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• Angiosperms began to replace gymnosperms
near the end of the Mesozoic
• Angiosperms now dominate more terrestrial
ecosystems
• Today, cone-bearing gymnosperms called
conifers dominate in the northern latitudes
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• The gymnosperm consist of four phyla
– Cycadophyta (cycads)
– Gingkophyta (one living species: Ginkgo biloba)
– Gnetophyta (three genera: Gnetum, Ephedra,
Welwitschia)
– Coniferophyta (conifers, such as pine, fir, and
redwood)
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Phylum Cycadophyta
• Individuals have large cones and palmlike leaves
• These thrived during the Mesozoic, but relatively
few species exist today
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Figure 30.5a
Cycas revoluta
Phylum Ginkgophyta
• This phylum consists of a single living species,
Ginkgo biloba
• It has a high tolerance to air pollution and is a
popular ornamental tree
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Ginkgo biloba leaves and fleshy seeds
Ginkgo biloba pollen-producing tree
Figure 30.5b
Figure 30.5ba
Ginkgo biloba pollen-producing tree
Figure 30.5bb
Ginkgo biloba leaves
and fleshy seeds
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Phylum Gnetophyta
• This phylum comprises three genera
• Species vary in appearance, and some are
tropical whereas others live in deserts
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Figure 30.5d
Ovulate cones Gnetum
Ephedra
Welwitschia
Figure 30.5da
Welwitschia
Figure 30.5db
Ovulate cones
Welwitschia
Figure 30.5dc
Gnetum
Figure 30.5dd
Ephedra
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Phylum Coniferophyta
• This phylum is by far the largest of the
gymnosperm phyla
• Most conifers are evergreens and can carry out
photosynthesis year round
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Figure 30.5e
Douglas fir
Common juniper
European larch
Sequoia
Wollemi pine Bristlecone pine
Figure 30.5ea
Douglas fir
Figure 30.5eb
European larch
Figure 30.5ec
Sequoia
Figure 30.5ed
Common juniper
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Figure 30.5ee
Wollemi pine
Figure 30.5ef
Wollemi pine
Figure 30.5eg
Bristlecone pine
The Life Cycle of a Pine: A Closer Look
• Three key features of the gymnosperm life cycle
are
– Dominance of the sporophyte generation
– Development of seeds from fertilized ovules
– The transfer of sperm to ovules by pollen
• The life cycle of a pine provides an example
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Animation: Pine Life Cycle
• The pine tree is the sporophyte and produces sporangia in male and female cones
• Small cones produce microspores called pollen grains, each of which contains a male gametophyte
• The familiar larger cones contain ovules, which produce megaspores that develop into female gametophytes
• It takes nearly three years from cone production to mature seed
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Key
Haploid (n)
Diploid (2n)
Mature sporophyte (2n)
Pollen cone
Microsporocytes (2n)
Microsporangia
Microsporangium (2n)
Pollen grains (n)
MEIOSIS
Figure 30.6-1
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Key
Haploid (n)
Diploid (2n)
Mature sporophyte (2n)
Ovulate cone
Pollen cone
Microsporocytes (2n)
Microsporangia
Microsporangium (2n) Surviving megaspore (n)
MEIOSIS
Megasporangium (2n) Pollen grain
Pollen grains (n)
MEIOSIS
Megasporocyte (2n)
Integument
Ovule
Figure 30.6-2
Key
Haploid (n)
Diploid (2n)
Mature sporophyte (2n)
Ovulate cone
Pollen cone
Microsporocytes (2n)
Microsporangia
Microsporangium (2n)
Archegonium
Surviving megaspore (n)
MEIOSIS
Megasporangium (2n) Pollen grain
Pollen grains (n)
MEIOSIS
Female gametophyte
Megasporocyte (2n)
Integument
Sperm nucleus (n) Egg nucleus (n)
Pollen tube
FERTILIZATION
Ovule
Figure 30.6-3
Key
Haploid (n)
Diploid (2n)
Mature sporophyte (2n)
Ovulate cone
Pollen cone
Microsporocytes (2n)
Microsporangia
Microsporangium (2n)
Seedling
Archegonium
Surviving megaspore (n)
MEIOSIS
Megasporangium (2n) Pollen grain
Pollen grains (n)
MEIOSIS
Female gametophyte
Megasporocyte (2n)
Integument
Sperm nucleus (n) Egg nucleus (n)
Pollen tube
Seed coat (2n)
FERTILIZATION
Food reserves (n)
Seeds
Embryo (new sporophyte) (2n)
Ovule
Figure 30.6-4
Concept 30.3: 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
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Figure 30.UN02
Nonvascular plants (bryophytes)
Seedless vascular plants
Gymnosperms
Angiosperms
Characteristics of Angiosperms
• All angiosperms are classified in a single phylum,
Anthophyta, from the Greek anthos for flower
• Angiosperms have two key adaptations
– Flowers
– Fruits
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Flowers
• The flower is an angiosperm structure
specialized for sexual reproduction
• Many species are pollinated by insects or
animals, while some species are wind-pollinated
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• A flower is a specialized shoot with up to four
types of modified leaves
– Sepals, which enclose the flower
– Petals, which are brightly colored and attract
pollinators
– Stamens, which produce pollen
– Carpels, which produce ovules
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• A stamen consists of a stalk called a filament,
with a sac called an anther where the pollen is
produced
• A carpel consists of an ovary at the base and a
style leading up to a stigma, where pollen is
received
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Video: Flower Blooming (time lapse)
Stamen Anther
Filament
Stigma Carpel
Style
Ovary
Petal
Sepal
Ovule
Figure 30.7
Fruits
• A fruit typically consists of a mature ovary but
can also include other flower parts
• Fruits protect seeds and aid in their dispersal
• Mature fruits can be either fleshy or dry
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Animation: Fruit Development
Tomato Ruby grapefruit
Hazelnut
Nectarine
Milkweed
Figure 30.8
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Figure 30.8a
Tomato
Figure 30.8b
Ruby grapefruit
Figure 30.8c
Nectarine
Figure 30.8d
Hazelnut
Figure 30.8e
Milkweed
• Various fruit adaptations help disperse seeds
• Seeds can be carried by wind, water, or animals
to new locations
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Wings
Seeds within berries
Barbs
Figure 30.9 Figure 30.9a
Wings
Figure 30.9b
Seeds within berries
Figure 30.9c
Barbs
Figure 30.9d
Barbs
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, or 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
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• A pollen grain that has landed on a stigma
germinates and the pollen tube of the male
gametophyte grows down to the ovary
• The ovule is entered by a pore called the
micropyle
• Double fertilization occurs when the pollen tube
discharges two sperm into the female
gametophyte within an ovule
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• One sperm fertilizes the egg, while the other
combines with two nuclei in the central cell of the
female gametophyte and initiates development of
food-storing endosperm
• The triploid endosperm nourishes the developing
embryo
• Within a seed, the embryo consists of a root and
two seed leaves called cotyledons
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Anther Mature flower on sporophyte plant (2n)
Pollen grains
Tube cell
Generative cell
Microspore (n) MEIOSIS
Microsporangium
Microsporocytes (2n)
Key
Haploid (n)
Diploid (2n)
Figure 30.10-1
Anther Mature flower on sporophyte plant (2n)
Megasporangium (2n)
Ovary
Antipodal cells
Central cell
Synergids
Female gametophyte
(embryo sac) Egg (n)
Surviving megaspore (n)
Pollen tube
Sperm (n)
Pollen grains
Tube cell
Generative cell
Microspore (n)
Male gametophyte
(in pollen grain) (n)
Ovule (2n)
MEIOSIS
MEIOSIS
Microsporangium
Microsporocytes (2n)
Key
Haploid (n)
Diploid (2n)
Figure 30.10-2
Anther Mature flower on sporophyte plant (2n)
Megasporangium (2n)
Ovary
Seed
Antipodal cells
Central cell
Synergids
Female gametophyte
(embryo sac) Egg (n)
Egg nucleus (n)
Surviving megaspore (n)
Pollen tube
Sperm (n)
Style
Sperm Pollen tube
Stigma
Pollen grains
Tube cell
Generative cell
Microspore (n)
Male gametophyte
(in pollen grain) (n)
Ovule (2n)
MEIOSIS
MEIOSIS
Discharged sperm nuclei (n)
FERTILIZATION
Microsporangium
Microsporocytes (2n)
Key
Haploid (n)
Diploid (2n)
Figure 30.10-3
Anther Mature flower on sporophyte plant (2n)
Germinating seed
Megasporangium (2n)
Ovary
Embryo (2n)
Endosperm (3n)
Seed coat (2n)
Seed
Antipodal cells
Central cell
Synergids
Female gametophyte
(embryo sac) Egg (n)
Egg nucleus (n)
Surviving megaspore (n)
Pollen tube
Sperm (n)
Style
Sperm Pollen tube
Stigma
Pollen grains
Tube cell
Generative cell
Microspore (n)
Male gametophyte
(in pollen grain) (n)
Ovule (2n)
MEIOSIS
MEIOSIS
Discharged sperm nuclei (n)
FERTILIZATION Zygote (2n)
Microsporangium
Microsporocytes (2n)
Nucleus of developing endosperm (3n)
Key
Haploid (n)
Diploid (2n)
Figure 30.10-4
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Video: Flowering Plant Life Cycle (time lapse)
Animation: Seed Development
Animation: Plant Fertilization
Angiosperm Evolution
• Darwin called the origin of angiosperms an
“abominable mystery”
• Angiosperms originated at least 140 million years
ago
• During the late Mesozoic, the major branches of
the clade diverged from their common ancestor
• Scientists are studying fossils, refining
phylogenies, and investigating developmental
patterns to resolve the mystery
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Fossil Angiosperms
• Chinese fossils of 125-million-year-old
angiosperms share some traits with living
angiosperms but lack others
• Archaefructus sinensis, for example, has anthers
and seeds but lacks petals and sepals
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(a) Archaefructus sinensis, a 125- million-year-old fossil
(b) Artist’s reconstruction of Archaefructus sinensis
Carpel
Stamen
5 cm
Figure 30.11
Figure 30.11a
(a) Archaefructus sinensis, a 125- million-year-old fossil
5 cm
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
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Microsporangia
(contain
microspores)
Ovules
Most recent common ancestor
of all living angiosperms
300 250 200 150 100 50 0
Eudicots
Monocots
Magnoliids
Star anise
and relatives
Water lilies
Amborella
Bennettitales
Living
gymnosperms
Millions of years ago
(b) Angiosperm phylogeny
(a) A possible ancestor of the angiosperms?
Figure 30.12
Microsporangia (contain microspores)
Ovules
(a) A possible ancestor of the angiosperms?
Figure 30.12a
Most recent common ancestor of all living angiosperms
300
Eudicots
Monocots
Magnoliids
Star anise
and relatives
Water lilies
Amborella
Bennettitales
Living
gymnosperms
Millions of years ago
(b) Angiosperm phylogeny
250 200 150 100 50 0
Figure 30.12b
Developmental Patterns in Angiosperms
• Egg formation in the angiosperm Amborella
resembles that of the gymnosperms
• In early angiosperms, the two integuments
appear to originate separately
• Researchers are currently studying expression of
flower development genes in gymnosperm and
angiosperm species
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Angiosperm Diversity
• Angiosperms comprise more than 250,000 living species
• Previously, angiosperms were divided into two main groups
– Monocots (one cotyledon)
– Dicots (two dicots)
• DNA studies suggest that monocots form a clade, but dicots are polyphyletic
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• The clade eudicot (“true” dicots) includes most
dicots
• The rest of the former dicots form several small
lineages
• Basal angiosperms are less derived and
include the flowering plants belonging to the
oldest lineages
• Magnoliids share some traits with basal
angiosperms but evolved later
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Basal Angiosperms
• Three small lineages constitute the basal
angiosperms
• These include Amborella trichopoda, water lilies,
and star anise
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Water lily
Star anise
Amborella trichopoda
Basal Angiosperms Figure 30.13a
Water lily
Figure 30.13aa Figure 30.13ab
Star anise
Figure 30.13ac
Amborella trichopoda
Magnoliids
• Magnoliids include magnolias, laurels, and black
pepper plants
• Magnoliids are more closely related to monocots
and eudicots than basal angiosperms
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Figure 30.13b
Magnoliids
Southern magnolia
Monocots
• More than one-quarter of angiosperm species
are monocots
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Orchid
Monocots
Lily
Pygmy date palm
Anther
Filament
Stigma
Ovary
Barley, a grass
Figure 30.13c Figure 30.13ca
Orchid
Figure 30.13cb
Pygmy date palm
Figure 30.13cc
Lily
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Figure 30.13cd
Anther
Filament
Stigma
Ovary
Barley, a grass
Barley, a grass
Figure 30.13ce
Eudicots
• More than two-thirds of angiosperm species are
eudicots
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California poppy Dog rose
Pyrenean oak
Snow pea Zucchini
Eudicots Figure 30.13d
Figure 30.13da
California poppy
Figure 30.13db
Pyrenean oak
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Figure 30.13dc
Dog rose
Figure 30.13dd
Snow pea
Figure 30.13de
Zucchini
Figure 30.13e Monocot
Characteristics Eudicot
Characteristics
Embryos
One cotyledon Two cotyledons
Leaf venation
Veins usually parallel
Veins usually netlike
Stems
Vascular tissue scattered
Vascular tissue usually arranged
in ring
Roots
Root system usually fibrous
(no main root)
Taproot (main root) usually present
Pollen
Pollen grain with one opening
Pollen grain with three openings
Flowers
Floral organs usually in
multiples of three
Floral organs usually in multiples
of four or five
Figure 30.13ea
Monocot
Characteristics
Eudicot
Characteristics
Embryos
One cotyledon Two cotyledons
Leaf
venation
Veins usually
parallel
Veins usually
netlike
Stems
Vascular tissue
scattered
Vascular tissue
usually arranged
in ring
Figure 30.13eb
Roots
Root system
usually fibrous
(no main root)
Taproot (main root)
usually present
Pollen
Pollen grain with
one opening Pollen grain with
three openings
Flowers
Floral organs
usually in
multiples of three
Floral organs
usually in multiples
of four or five
Monocot Characteristics
Eudicot Characteristics
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Evolutionary Links Between Angiosperms
and Animals
• Animals influence the evolution of plants and vice
versa
– For example, animal herbivory selects for plant
defenses
– For example, interactions between pollinators
and flowering plants select for mutually beneficial
adaptations
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Video: Bee Pollinating
Video: Bat Pollinating Agave Plant
Figure 30.14
• 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
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Figure 30.15
Concept 30.4: 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
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Products from Seed Plants
• Most of our food comes from angiosperms
• Six crops (wheat, rice, maize, potatoes, cassava, and sweet potatoes) yield 80% of the calories consumed by humans
• Modern crops are products of relatively recent genetic change resulting from artificial selection
• Many seed plants provide wood
• Secondary compounds of seed plants are used in medicines
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Table 30.1
Threats to Plant Diversity
• Destruction of habitat is causing extinction of
many plant species
• In the tropics 55,000 km2 are cleared each year
• At this rate, the remaining tropical forests will be
eliminated in 200 years
• Loss of plant habitat is often accompanied by
loss of the animal species that plants support
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• At the current rate of habitat loss, 50% of Earth’s
species will become extinct within the next 100–
200 years
• The tropical rain forests may contain
undiscovered medicinal compounds
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Figure 30.16
A satellite image from 2000 shows clear-cut areas in Brazil surrounded by dense tropical forest.
By 2009, much more of this same tropical forest had been cut down.
4 k
m
Figure 30.16a
A satellite image
from 2000 shows
clear-cut areas in
Brazil surrounded by
dense tropical forest.
4 k
m
Figure 30.16b
4 k
m
By 2009, much more of this same tropical forest had been cut down.
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Figure 30.UN03
Time since divergence from common ancestor
Common ancestor “Bilateral” clade
“Radial” clade
Compare numbers of species
Five Derived Traits of Seed Plants
Reduced gametophytes
Heterospory
Ovules
Pollen
Seeds
Microscopic male and female gametophytes (n) are nourished and protected by the sporophyte (2n)
Microspore (gives rise to a male gametophyte)
Megaspore (gives rise to a female gametophyte)
Ovule (gymnosperm)
Integument (2n)
Megaspore (n)
Megasporangium (2n)
Pollen grains make water unnecessary for fertilization
Seeds: survive better than unprotected spores, can be transported long distances
Seed coat
Food supply
Embryo
Female gametophyte
Male gametophyte
Figure 30.UN04
Figure 30.UN05
Charophyte green algae
Mosses
Ferns
Gymnosperms
Angiosperms
Figure 30.UN06
Charophyte green algae
Mosses
Ferns
Gymnosperms
Angiosperms
Figure 30.UN07