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 38

Angiosperm Reproduction
and Biotechnology

Lectures by
Erin Barley
Kathleen Fitzpatrick

© 2011 Pearson Education, Inc.

Overview: Flowers of Deceit
• Insects help angiosperms to reproduce sexually
with distant members of their own species
– For example, male Campsoscolia wasps mistake
Ophrys flowers for females and attempt to mate
with them
– The flower is pollinated in the process
– Unusually, the flower does not produce nectar
and the male receives no benefit

© 2011 Pearson
Copyright © 2008Education, Inc.
Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Figure 38.1
 • Many angiosperms lure insects with nectar; both
plant and pollinator benefit
• Mutualistic symbioses are common between
plants and other species
• Angiosperms can reproduce sexually and
asexually
• Angiosperms are the most important group of
plants in terrestrial ecosystems and in agriculture

© 2011 Pearson
Copyright © 2008Education, Inc.
Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Concept 38.1: Flowers, double fertilization,
and fruits are unique features of the
angiosperm life cycle
• Plant lifecycles are characterized by the alternation
between a multicellular haploid (n) generation and
a multicellular diploid (2n) generation
• Diploid sporophytes (2n) produce spores (n) by
meiosis; these grow into haploid gametophytes (n)
• Gametophytes produce haploid gametes (n) by
mitosis; fertilization of gametes produces a
sporophyte

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 • In angiosperms, the sporophyte is the dominant
generation, the large plant that we see
• The gametophytes are reduced in size and
depend on the sporophyte for nutrients
• The angiosperm life cycle is characterized by
“three Fs”: flowers, double fertilization, and fruits

© 2011 Pearson Education, Inc.

Figure 38.2

Anther Germinated pollen grain (n)

Stamen Stigma Carpel
Anther (male gametophyte)
Style Ovary
Filament Ovary Pollen tube
Ovule
Embryo sac (n)
(female gametophyte)

Sepal FERTILIZATION
Petal Egg (n)
Sperm (n)
Receptacle
Mature sporophyte Zygote
(a) Structure of an Key plant (2n) (2n)
idealized flower Seed
Haploid (n) Germinating
Diploid (2n) seed
Seed

Embryo (2n)
(b) Simplified angiosperm Simple (sporophyte)
life cycle fruit
Flower Structure and Function
• Flowers are the reproductive shoots of the
angiosperm sporophyte; they attach to a part of
the stem called the receptacle
• Flowers consist of four floral organs: sepals,
petals, stamens, and carpels
• Stamens and carpels are reproductive organs;
sepals and petals are sterile

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 • A stamen consists of a filament topped by an
anther with pollen sacs that produce pollen
• A carpel has a long style with a stigma on which
pollen may land
• At the base of the style is an ovary containing one
or more ovules
• A single carpel or group of fused carpels is called
a pistil

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 • Complete flowers contain all four floral organs
• Incomplete flowers lack one or more floral
organs, for example stamens or carpels
• Clusters of flowers are called inflorescences

© 2011 Pearson Education, Inc.

Development of Male Gametophytes in
Pollen Grains
• Pollen develops from microspores within
the microsporangia, or pollen sacs, of anthers
• Each microspore undergoes mitosis to produce
two cells: the generative cell and the tube cell
• A pollen grain consists of the two-celled male
gametophyte and the spore wall

© 2011 Pearson Education, Inc.

 • If pollination succeeds, a pollen grain produces a
pollen tube that grows down into the ovary and
discharges two sperm cells near the embryo sac

© 2011 Pearson Education, Inc.

Figure 38.3
(a) Development of a male (b) Development of a female
gametophyte gametophyte (embryo sac)
(in pollen grain)

Microsporangium
(pollen sac) Megasporangium

Microsporocyte Ovule Megasporocyte

MEIOSIS
Integuments
Microspores (4) Micropyle

Surviving
megaspore
Each of 4
microspores
MITOSIS

Female gametophyte
Male Ovule Antipodal cells (3)
Generative cell

(embryo sac)
(will form 2 gametophyte
sperm) (in pollen grain) Polar nuclei (2)

Egg (1)
Nucleus of tube cell
Integuments
Synergids (2)
20 m
Ragweed
pollen Key to labels
grain Embryo sac
100 m

75 m (colorized Haploid (n)

(LM) SEM) Diploid (2n) (LM)
Development of Female Gametophytes
(Embryo Sacs)
• The embryo sac, or female gametophyte,
develops within the ovule
• Within an ovule, two integuments surround a
megasporangium
• One cell in the megasporangium undergoes
meiosis, producing four megaspores, only one of
which survives
• The megaspore divides, producing a large cell
with eight nuclei

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 • This cell is partitioned into a multicellular female
gametophyte, the embryo sac

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Pollination
• In angiosperms, pollination is the transfer of
pollen from an anther to a stigma
• Pollination can be by wind, water, or animals
• Wind-pollinated species (e.g., grasses and many
trees) release large amounts of pollen

© 2011 Pearson Education, Inc.

Figure 38.4a
Abiotic Pollination by Wind Pollination by Bees

Common dandelion
under normal light

Hazel staminate flowers

(stamens only)
Hazel carpellate
flower (carpels only)

Common dandelion
under ultraviolet light
Figure 38.4b
Pollination by Moths
and Butterflies Pollination by Flies Pollination by Bats

Anther
Moth

Fly egg

Stigma
Blowfly on carrion Long-nosed bat feeding
Moth on yucca flower flower on cactus flower at night

Pollination by Birds

Hummingbird
drinking nectar of
columbine flower
Coevolution of Flower and Pollinator
• Coevolution is the evolution of interacting species
in response to changes in each other
• Many flowering plants have coevolved with
specific pollinators
• The shapes and sizes of flowers often correspond
to the pollen transporting parts of their animal
pollinators
– For example, Darwin correctly predicted a moth
with a 28 cm long tongue based on the
morphology of a particular flower

© 2011 Pearson Education, Inc.

Figure 38.5
Double Fertilization
• After landing on a receptive stigma, a pollen grain
produces a pollen tube that extends between the
cells of the style toward the ovary
• Double fertilization results from the discharge of
two sperm from the pollen tube into the embryo sac
• One sperm fertilizes the egg, and the other combines
with the polar nuclei, giving rise to the triploid food-
storing Plant Fertilization endosperm (3n)

© 2011 Pearson Education, Inc.

 Animation: Plant Fertilization
Right-click slide / select “Play”
© 2011 Pearson Education, Inc.
Figure 38.6-3

1 2 3
Stigma Pollen Endosperm
grain nucleus (3n)
Pollen (2 polar nuclei
tube Ovule plus sperm)
2 sperm Polar
nuclei
Style
Ovary Egg
Zygote
Ovule Synergid (2n)
Polar
nuclei 2 sperm
Egg
Micropyle
Seed Development, Form, and Function
• After double fertilization, each ovule develops into
a seed
• The ovary develops into a fruit enclosing the
seed(s)

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Endosperm Development
• Endosperm development usually precedes
embryo development
• In most monocots and some eudicots, endosperm
stores nutrients that can be used by the seedling
• In other eudicots, the food reserves of the
endosperm are exported to the cotyledons

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Embryo Development
• The first mitotic division of the zygote splits the
fertilized egg into a basal cell and a terminal cell
• The basal cell produces a multicellular suspensor,
which anchors the embryo to the parent plant
• The terminal cell gives rise to most of the embryo
• The cotyledons form and the embryo elongates

© 2011 Pearson Education, Inc.

 Animation: Seed Development
Right-click slide / select “Play”
© 2011 Pearson Education, Inc.
Figure 38.7

Ovule
Proembryo
Endosperm
nucleus Suspensor
Integuments
Cotyledons
Zygote Basal Shoot
cell apex
Root
apex
Seed
Suspensor coat
Terminal cell
Basal cell Endosperm
Zygote
Structure of the Mature Seed
• The embryo and its food supply are enclosed by a
hard, protective seed coat
• The seed enters a state of dormancy
• A mature seed is only about 5–15% water

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 • In some eudicots, such as the common garden
bean, the embryo consists of the embryonic axis
attached to two thick cotyledons (seed leaves)
• Below the cotyledons the embryonic axis is called
the hypocotyl and terminates in the radicle
(embryonic root); above the cotyledons it is called
the epicotyl
• The plumule comprises the epicotyl, young leaves,
and shoot apical meristem

© 2011 Pearson Education, Inc.

Figure 38.8
Seed coat Epicotyl
Hypocotyl

Radicle
Cotyledons

(a) Common garden bean, a eudicot with thick cotyledons

Seed coat
Endosperm
Cotyledons
Epicotyl
Hypocotyl
Radicle

(b) Castor bean, a eudicot with thin cotyledons

Scutellum Pericarp fused

(cotyledon) with seed coat
Endosperm
Coleoptile
Epicotyl
Hypocotyl
Coleorhiza
Radicle
(c) Maize, a monocot
 • The seeds of some eudicots, such as castor
beans, have thin cotyledons

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 • A monocot embryo has one cotyledon
• Grasses, such as maize and wheat, have a
special cotyledon called a scutellum
• Two sheathes enclose the embryo of a grass
seed: a coleoptile covering the young shoot and a
coleorhiza covering the young root

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Seed Dormancy: An Adaptation for Tough
Times
• Seed dormancy increases the chances that
germination will occur at a time and place most
advantageous to the seedling
• The breaking of seed dormancy often requires
environmental cues, such as temperature or
lighting changes

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Seed Germination and Seedling Development
• Germination depends on imbibition, the uptake of
water due to low water potential of the dry seed
• The radicle (embryonic root) emerges first
• Next, the shoot tip breaks through the soil surface

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 • In many eudicots, a hook forms in the hypocotyl,
and growth pushes the hook above ground
• Light causes the hook to straighten and pull the
cotyledons and shoot tip up

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Figure 38.9
Foliage leaves
Cotyledon
Epicotyl
Hypocotyl
Cotyledon Cotyledon
Hypocotyl
Hypocotyl

Radicle
Seed coat
(a) Common garden bean

Foliage leaves

Coleoptile Coleoptile

Radicle
(b) Maize
 • In maize and other grasses, which are monocots,
the coleoptile pushes up through the soil

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Fruit Form and Function
• A fruit develops from the ovary
• It protects the enclosed seeds and aids in seed
dispersal by wind or animals
• A fruit may be classified as dry, if the ovary dries
out at maturity, or fleshy, if the ovary becomes
thick, soft, and sweet at maturity

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 Animation: Fruit Development
Right-click slide / select “Play”
© 2011 Pearson Education, Inc.
 • Fruits are also classified by their development
– Simple, a single or several fused carpels
– Aggregate, a single flower with multiple separate
carpels
– Multiple, a group of flowers called an
inflorescence

© 2011 Pearson Education, Inc.

Figure 38.10

Stigma Style
Carpels Stamen Flower Petal
Stamen Ovary

Stamen
Sepal
Stigma Ovary (in
Ovule receptacle)
Ovule
Pea flower Raspberry flower Pineapple Apple flower
inflorescence
Each segment Remains of
Carpel stamens and styles
develops
(fruitlet)
Stigma from the Sepals
Seed
Ovary carpel
of one
Stamen flower

Seed
Receptacle
Pea fruit Raspberry fruit Pineapple fruit Apple fruit
(a) Simple fruit (b) Aggregate fruit (c) Multiple fruit (d) Accessory fruit
 • An accessory fruit contains other floral parts in
addition to ovaries

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 • Fruit dispersal mechanisms include
– Water
– Wind
– Animals

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Figure 38.11a
Dispersal by Wind

Dandelion fruit
Dandelion “seeds” (actually one-seeded fruits) Tumbleweed

Winged fruit of a maple Winged seed of

the tropical Asian
climbing gourd
Alsomitra macrocarpa

Dispersal by Water

Coconut seed embryo,

endosperm, and endocarp
inside buoyant husk
Figure 38.11b

Dispersal by Animals
Fruit of puncture vine
(Tribulus terrestris)

Squirrel hoarding
seeds or fruits
underground

Ant carrying
seed with nutritious
“food body” to its
nest

Seeds dispersed in black bear feces

Concept 38.2: Flowering plants reproduce
sexually, asexually, or both
• Many angiosperm species reproduce both
asexually and sexually
• Sexual reproduction results in offspring that are
genetically different from their parents
• Asexual reproduction results in a clone of
genetically identical organisms

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Mechanisms of Asexual Reproduction
• Fragmentation, separation of a parent plant into
parts that develop into whole plants, is a very
common type of asexual reproduction
• In some species, a parent plant’s root system
gives rise to adventitious shoots that become
separate shoot systems

© 2011 Pearson Education, Inc.

Figure 38.12
 • Apomixis is the asexual production of seeds from
a diploid cell

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Advantages and Disadvantages of Asexual
Versus Sexual Reproduction
• Asexual reproduction is also called vegetative
reproduction
• Asexual reproduction can be beneficial to a
successful plant in a stable environment
• However, a clone of plants is vulnerable to local
extinction if there is an environmental change

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 • Sexual reproduction generates genetic variation
that makes evolutionary adaptation possible
• However, only a fraction of seedlings survive
• Some flowers can self-fertilize to ensure that every
ovule will develop into a seed
• Many species have evolved mechanisms to
prevent selfing

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Mechanisms That Prevent Self-Fertilization
• Many angiosperms have mechanisms that make it
difficult or impossible for a flower to self-fertilize
• Dioecious species have staminate and carpellate
flowers on separate plants

© 2011 Pearson Education, Inc.

Figure 38.13

(a) Staminate flowers (left) and carpellate flowers (right)

of a dioecious species

Stamens Styles
Styles Stamens

Thrum flower Pin flower

(b) Thrum and pin flowers
 • The most common is self-incompatibility, a
plant’s ability to reject its own pollen
• Researchers are unraveling the molecular
mechanisms involved in self-incompatibility
• Some plants reject pollen that has an S-gene
matching an allele in the stigma cells
• Recognition of self pollen triggers a signal
transduction pathway leading to a block in growth
of a pollen tube

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Vegetative Propagation and Agriculture
• Humans have devised methods for asexual
propagation of angiosperms
• Most methods are based on the ability of plants to
form adventitious roots or shoots

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Clones from Cuttings
• Many kinds of plants are asexually reproduced
from plant fragments called cuttings
• A callus is a mass of dividing undifferentiated
cells that forms where a stem is cut and produces
adventitious roots

© 2011 Pearson
Copyright © 2008Education, Inc.
Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Grafting
• A twig or bud can be grafted onto a plant of a
closely related species or variety
• The stock provides the root system
• The scion is grafted onto the stock

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Test-Tube Cloning and Related Techniques
• Plant biologists have adopted in vitro methods to
create and clone novel plant varieties
• A callus of undifferentiated cells can sprout shoots
and roots in response to plant hormones

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Figure 38.14

(a) (b) (c) Developing root

 • Transgenic plants are genetically modified (GM)
to express a gene from another organism
• Protoplast fusion is used to create hybrid plants
by fusing protoplasts, plant cells with their cell
walls removed

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Figure 38.15

50 m
Concept 38.3: Humans modify crops by
breeding and genetic engineering
• Humans have intervened in the reproduction and
genetic makeup of plants for thousands of years
• Hybridization is common in nature and has been
used by breeders to introduce new genes
• Maize, a product of artificial selection, is a staple
in many developing countries

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Figure 38.16
Plant Breeding
• Mutations can arise spontaneously or can be
induced by breeders
• Plants with beneficial mutations are used in
breeding experiments
• Desirable traits can be introduced from different
species or genera
• The grain triticale is derived from a successful
cross between wheat and rye

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Plant Biotechnology and Genetic
Engineering
• Plant biotechnology has two meanings
– In a general sense, it refers to innovations in the
use of plants to make useful products
– In a specific sense, it refers to use of GM
organisms in agriculture and industry
• Modern plant biotechnology is not limited to
transfer of genes between closely related species
or varieties of the same species

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Reducing World Hunger and Malnutrition
• Genetically modified plants may increase the
quality and quantity of food worldwide
• Transgenic crops have been developed that
– Produce proteins to defend them against insect
pests
– Tolerate herbicides
– Resist specific diseases

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 • Nutritional quality of plants is being improved
– For example, “Golden Rice” is a transgenic variety
being developed to address vitamin A deficiencies
among the world’s poor

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Figure 38.17

Cassava roots harvested in Thailand

Reducing Fossil Fuel Dependency
• Biofuels are made by the fermentation and
distillation of plant materials such as cellulose
• Biofuels can be produced by rapidly growing crops
such as switchgrass and poplar
• Biofuels would reduce the net emission of CO 2, a
greenhouse gas
• The environmental implications of biofuels are
controversial

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The Debate over Plant Biotechnology
• Some biologists are concerned about risks of
releasing GM organisms (GMOs) into the
environment

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Issues of Human Health
• One concern is that genetic engineering may
transfer allergens from a gene source to a plant
used for food
• Some GMOs have health benefits
– For example, maize that produces the Bt toxin
has 90% less of a cancer-causing toxin than non-
Bt corn
– Bt maize has less insect damage and lower
infection by Fusarium fungus that produces the
cancer-causing toxin

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 • GMO opponents advocate for clear labeling of all
GMO foods

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Possible Effects on Nontarget Organisms
• Many ecologists are concerned that the growing of
GM crops might have unforeseen effects on
nontarget organisms

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Addressing the Problem of Transgene Escape
• Perhaps the most serious concern is the
possibility of introduced genes escaping into
related weeds through crop-to-weed hybridization
• This could result in “superweeds” that would be
resistant to many herbicides

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 • Efforts are underway to prevent this by
introducing
– Male sterility
– Apomixis
– Transgenes into chloroplast DNA (not
transferred by pollen)
– Strict self-pollination

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