The Evolution of Seed Plants

KEY CONCEPTS
27
27.1 Pollen, Seeds, and Wood
Contributed to the Success of
Seed Plants
27.2 Once Dominant Gymnosperms
Still Thrive in Some
Environments
27.3 Flowers and Fruits Led to
Increased Diversification of
Angiosperms
27.4 Plants Play Critical Roles in
Terrestrial Ecosystems

© Zvonimir Atletić/Alamy Stock Photo

Phoenix dactylifera, the date palm, is widely

cultivated for its edible fruit.

▶InvestigatingLIFE

Brought Back from Extinction by a Seed

The Judean date is a fruit that was once much prized, admired environmental extremes through what may be a very long and
for its nutritional and medicinal properties. This fruit was the stressful resting period—in the case of the Judean date, many
source of the “honey” in the biblical “land of milk and honey.” centuries in a harsh desert. Such hardiness has contributed
Today that ancient strain of date is gone. Or is it? to making seed plants the predominant plants on Earth. All of
About 2,000 years ago, a seed developed in a fruit on a today’s forests are dominated by seed plants.
Judean date palm. The fruit that contained that seed found its So will the resurrected palm serve as the parent of a new
way to a storeroom in the fortress Masada in Judea. In 73 CE a population of Judean dates, thus bringing that genotype back
band of Jewish zealots involved in a revolt against Rome fled to from extinction? Unfortunately, it cannot do so alone, because
this refuge with their families and provisions. The fortress was date palms have separate sexes, and the single known indi-
eventually sacked and left in ruins. vidual is a male that can produce only pollen. However, the
Twenty centuries later, archeologists working in Masada dis- search is on for other ancient seeds of Judean dates from
covered the date seed that was long ago stored in the Masada archeological sites, and it is hoped that one of those will grow
fortress and confirmed its age. The previous record for seed sur- into a female plant that can produce new fruit and seeds. Even
vival and germination was 1,300 years, held by lotus seeds that if a female Judean date seed cannot be found, the male Judean
recently germinated under the care of scientists in China. But date’s pollen can be used to introduce valuable genes back into
botanist Elaine Solowey succeeded in making the 2,000-year-old other, still living, strains of dates.
date seed germinate! For 10 years the seedling has thrived, and
is now mature and producing pollen.
Seeds are important structures for the evolutionary survival
of plants. Each seed protects an enclosed plant embryo from
QA
and
How long can most seeds survive, and
why is seed dormancy important?
(Find the answer on page 620.)

27_Life12e_Ch 27.indd 1 12/4/19 9:42 AM

 CHAPTER 27 The Evolution of Seed Plants

species of which are now extinct. The progymnosperms included

KEY CONCEPT Pollen, Seeds, and Wood many large trees.
Contributed to the Success
27.1 of Seed Plants
Two other innovations, pollen and seeds, arose in the seed
plants. Pollen increases reproductive opportunities in a terrestrial
environment, and seeds provide a secure and lasting structure that
Learning Objectives protects the dormant stage of the embryo. A plant embryo may
safely wait within its seed (in some cases for many years, or even
27.1.1 Explain how the evolution of pollen enabled seed
plants to thrive in terrestrial environments. centuries) until conditions are right for germination.
The earliest fossil evidence of seed plants is found in late De-
27.1.2 List three major advantages of having seeds.
vonian rocks. Like the progymnosperms, these now-extinct seed
27.1.3 Describe the earliest group of woody plants and list ferns were woody. They possessed fernlike foliage but had seeds
two advantages of woody plants.
attached to their leaves. By the end of the Permian, other groups
of seed plants had become dominant (Figure 27.1). Today’s living
By the late Devonian, more than 360 million years ago, Earth seed plants fall into two major groups, the gymnosperms (such
was home to a great variety of land plants, many of which we dis- as pines and cycads) and the hugely diverse group known as the
cussed in Chapter 26. The land plants shared the hot, humid ter- angiosperms (flowering plants).
restrial environment with insects, spiders, centipedes, and early
tetrapods. These plants and animals evolved together, each acting Features of the seed plant life cycle protect
as agents of natural selection on the other. In the Devonian, a gametes and embryos
new innovation appeared when some plants developed extensively In Key Concept 26.2 we described a major trend in land plant
thickened woody stems. Among the first plants with this adapta- evolution: the sporophyte became less dependent on the game-
tion were seedless vascular plants called progymnosperms, all tophyte, which became smaller in relation to the sporophyte.

Rise of Gymnosperms Angiosperms

seed plants dominant dominant
© iStock.com/Irina Tischenko

Shutterstock.com
© Dole/

Angiosperms (flowering plants)

Conifers
Shutterstock.com
© hypnotype/

Ginkgos
David McIntyre

Cycads

Seed ferns

Progymnosperms (seedless vascular trees with roots and leaves)

Quaternary
Rhyniophytes (seedless vascular plants without roots or leaves) Neogene

Ordovician Silurian Devonian Carboniferous Permian Triassic Jurassic Cretaceous Paleogene

Paleozoic Mesozoic Cenozoic

500 400 300 200 100 Present

Time (mya)

Figure 27.1 The Fossil Record of Seed Plant Evolution Carboniferous, but the earliest known fossils of flowering plants are
Woody growth evolved in the seedless progymnosperms. The now- from near the Jurassic–Cretaceous boundary. The flowering plants have
extinct seed ferns had woody growth, fernlike foliage, and seeds dominated most terrestrial environments through the Cenozoic. (mya,
attached to their leaves. New lineages of seed plants arose during the millions of years ago.)

27_Life12e_Ch 27.indd 2 12/4/19 9:42 AM

 KEY CONCEPT 27.1 Pollen, Seeds, and Wood Contributed to the Success of Seed Plants

Sporophyte (2n) Male gametophyte

(microgametophytes; n)

Sporophyte (2n)
Anther
The moss
gametophyte
nourishes the
sporophyte. Ovary

Flower
Female gametophyte
(megagametophyte; n)

The seed plant sporophyte

nourishes the developing
gametophytes.
Gametophyte (n)
Gametophyte (n)

The large sporophytes and the small

gametophytes of ferns are nutritionally
independent of one another.

Figure 27.2 The Relationship between Sporophyte and (brown) has been reduced and the sporophyte (blue) has become
Gametophyte In the course of plant evolution, the gametophyte more prominent.

Corylus avellana
This trend continued with the seed plants, whose gametophyte
generation is reduced even further than it is in the ferns (Figure
27.2). The haploid seed plant gametophyte develops partly or
entirely while attached to and nutritionally dependent on the
diploid sporophyte.
Among the seed plants, only the earliest-diverging groups of
gymnosperms (including modern cycads and ginkgos) have swim-
ming sperm. Even in these groups, sperm is transferred via pollen
grains, so fertilization does not require liquid water outside the
plant body. The evolution of pollen, along with the advent of seeds,
gave seed plants the opportunity to colonize drier areas and spread
over the terrestrial environment.
Seed plants are heterosporous (see Figure 26.18B)—that is, they
produce two types of spores, one that becomes a male gameto-
© Wildlife GmbH/Alamy Stock Photo

phyte (microgametophyte) and one that becomes a female game-

tophyte (macrogametophyte). They form separate microsporangia
and megasporangia on structures that are grouped on short stems,
such as the stamens and carpels of an angiosperm flower.
Within the microsporangium, the meiotic products are mi-
crospores. Within its spore wall, a microspore divides mitotically
one or a few times to form a multicellular male gametophyte,
also known as a single grain of pollen. Pollen grains are released Figure 27.3 Blown on the Wind Pollen grains are the male game-
from the microsporangium to be distributed by wind or by an tophytes of seed plants. The male flowers of this hazel tree release
animal pollinator (Figure 27.3). As in seedless land plants, the pollen-containing spores that are dispersed by the wind and may land
spore wall that surrounds the pollen grain contains a substance near female gametophytes of other hazel plants.
called sporopollenin, the most chemically resistant biological Media Clip 27.1 Pollen Transfer by Wind
compound known. Sporopollenin protects the pollen grain against Life12e.com/mc27.1

27_Life12e_Ch 27.indd 3 12/4/19 9:42 AM

 CHAPTER 27 The Evolution of Seed Plants

dehydration and chemical damage—another advantage in terms and digests its way toward the female gametophyte (Figure 27.4).
of survival in the terrestrial environment. When the tip of the pollen tube reaches the female gametophyte,
In contrast to the microspores, the megaspores of seed plants sperm are released from the tube and fertilization occurs.
are not shed. Instead, they develop into female gametophytes with- The resulting diploid zygote divides repeatedly, forming an em-
in the megasporangia. These female gametophytes are dependent bryonic sporophyte. After a period of embryonic development,
on the sporophyte for food and water. growth is temporarily suspended (the embryo enters a dormant
In most seed plant species, only one of the meiotic products stage). The end product at this stage is the multicellular seed.
in a megasporangium survives. The surviving haploid nucleus
divides mitotically, and the resulting cells divide again to pro- The seed is a complex, well-protected package
duce a multicellular female gametophyte. The megasporangium A seed contains tissues from three generations (Figure 27.5). A seed
is surrounded by sterile sporophytic structures, which form an coat develops from the integument—the tissues of the diploid sporo-
integument that protects the megasporangium and its contents. phyte parent that surround the megasporangium. Within the mega-
Together, the megasporangium and integument constitute the sporangium is haploid tissue from the female gametophyte, which
ovule, which will develop into a seed after fertilization. contains a supply of nutrients for the developing embryo. (This tis-
The arrival of a pollen grain at an appropriate landing point, sue is fairly extensive in most gymnosperm seeds. In angiosperm
close to a female gametophyte on a sporophyte of the same spe- seeds it is greatly reduced, and nutrition for the embryo is supplied
cies, is called pollination. A pollen grain that reaches this point instead by a tissue called endosperm.) In the center of the seed is
develops further. It produces a slender pollen tube that elongates the third generation, the embryo of the new diploid sporophyte.

(A) A growing pollen tube

Pollen grains
© Dr. Jeremy Burgess/Science Source

The pollen tube

elongates on its
way to the female
gametophyte.

(B) The process of pollination Pollen tube

The anthers of
Pollen grains Anther the stamen bear
pollen-producing
Filament microsporangia.
Petal

Stigma
The carpel Style
receives
pollen. Ovary
Ovule

When the tip of the

Sepal pollen tube reaches the
female gametophyte,
Integument
sperm are released from
Female gametophyte the tube and fertilization
ensues (see Figure 27.16).
Receptacle

Figure 27.4 Pollination Is a Hallmark of the Seed Plants In most growing in the flower of an angiosperm, the prairie gentian. (B) The pro-
seed plants, a pollen tube grows from the pollen grain to the female cess of pollination is diagrammed for a generalized angiosperm flower.
gametophyte, where sperm are released and fertilize the egg within the View in Achieve
ovule. Once fertilization takes place, the ovule can develop into a seed Activity 27.1 Flower Morphology
(see Figure 27.5). (A) Scanning electron micrograph of a pollen tube

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 KEY CONCEPT 27.1 Pollen, Seeds, and Wood Contributed to the Success of Seed Plants

(A) Unfertilized ovule (B) Fertilized ovule (C) Seed

1 The fleshy 2 The megaspore grows

Seed coat (derived from
megasporangium into a multicellular,
integument; parental
(2n) is protected by haploid female
sporophyte tissue)
the integument. gametophyte (n).

Immature female
pine cone
(cross section)
Food supply (female
gametophyte tissue; n)
Integument
Egg
nucleus (n) Embryo (new
Megaspore (n) sporophyte; 2n)
Germinated
pollen grain (n)

Micropyle

Pollen grain (n) 3 A pollen grain (n) enters through 4 The germinated pollen grain releases
the micropyle and develops a a sperm nucleus, fertilizing the egg
pollen tube (germinates). nucleus and initiating seed formation.

Figure 27.5 A Seed Develops These cross sections diagram the megasporangium). (B) The mature megaspore is fertilized by a pollen
development of the ovule into a seed in a gymnosperm (Pinus sp.). grain that penetrates the integument, germinates (grows a pollen tube;
Angiosperm seed development has differences (e.g., angiosperm integu- see Figure 27.4A), and releases a sperm nucleus. (C) Fertilization initi-
ments have two layers rather than one, and the angiosperm embryo is ates production of a seed. A mature seed contains three generations:
nourished by specialized tissue called endosperm) but follows the same a diploid embryo (the new sporophyte), which is surrounded by haploid
principle (compare Figures 27.8 and 27.16). (A) The haploid mega- female gametophyte tissue that supplies nutrition, which is in turn sur-
spore is nourished by tissues of the parental sporophyte (the diploid rounded by the seed coat (diploid parental sporophyte tissue).

The seed is a well-protected resting stage. As we discussed in a sandy knoll. The seeds experienced normal temperature fluctua-
the opening of this chapter, the seeds of some species may remain tions for Michigan. At regular intervals ever since, other biologists
dormant but stay viable (capable of growth and development) for have excavated a bottle and checked the viability of the seeds it
many years, germinating only when conditions are favorable for contained. The seeds of most species remained viable for decades,
the growth of the sporophyte. During the dormant stage, the seed whereas others have remained viable for more than a century.
coat protects the embryo from excessive drying and may also pro-
tect it against potential predators that would otherwise consume A change in stem anatomy enabled seed plants to
the embryo and its nutrient reserves. Many seeds have structural grow to great heights
adaptations or are contained in fruits that promote their dispersal Fossils of the closest relatives of seed plants (progymnosperms) and
by wind or, more often, by animals. When the young sporophyte the earliest seed plants (seed ferns) are found in late Devonian rocks
resumes growth, it draws on the food reserves in the seed. The (see Figure 27.1). These plants had thickened woody stems, developed
possession of seeds is a major reason for the enormous evolution- through the proliferation of xylem. This type of growth, which in-
ary success of the seed plants, which are the dominant life forms creases the diameter of stems and roots in many modern seed plants,
of most modern terrestrial floras. is called secondary growth. Its product is secondary xylem, or wood.
The germination of the Judean date described in the opening The younger portion of the wood produced by secondary growth
story is an extreme example of seed dormancy. How do we know is well adapted for water transport, but older wood becomes clogged
how long most seeds remain viable? To find out, William J. Beal, with resins or other materials. Although no longer functional in
a biologist at Michigan State University, decided to begin an ex- transport, the older wood continues to provide support for the plant.
periment in 1879 that he could not hope to finish in his lifetime This support allows woody plants to grow taller than other plants
(Investigating Life: William Beal’s Seed Viability Study). He pre- around them and thus capture more light for photosynthesis.
pared 20 lots of seeds for long-term storage. Each lot consisted of Not all seed plants are woody. In the course of seed plant evo-
50 seeds from each of 23 species of plants. He mixed each lot of lution, many groups lost the woody growth habit; however, other
seeds with sand and placed the mixture in an uncapped bottle, advantageous attributes helped them become established in an
then buried all the bottles upside down (so they would stay dry) on astonishing variety of places.

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 CHAPTER 27 The Evolution of Seed Plants

▶InvestigatingLIFE William Beal’s Seed Viability Study

RESULTS▸ The table shows the number of germinating seeds (of

Experiment
Original Papers: W. J. Beal. 1884. The vitality of seeds. Proc Soc
Prom Agr Sci 5: 44–46. the original 50) from three of the species in years 50–100 of this
H. T. Darlington. 1941. The sixty-year period for Dr. Beal’s seed ongoing experiment.
viability. Am J Bot 28: 271–273.
Years after burial
A. Kivilaan and R. S. Bandurski. 1981. The one hundred–year
Species 50 60 70 80 90 100
period for Dr. Beal’s seed viability experiment. Am J Bot 68:
1290–1292. Oenothera biennis 19 12 7 5 0 0
(evening primrose)
F. W. Telewski and J. Zeevaart. 2002. The 120-year period for Dr.
Beal's seed viability study. Am J Bot 89: 1285–1288. Rumex crispus (curly 26 2 7 1 0 0
dock)
William Beal began an experiment in 1879 to measure the long-
Verbascum blattaria 31 34 37 35 10 21
term viability of seeds of several common plants. This ongoing (moth mullein)
experiment has been continued by biologists for well over a

CONCLUSION▸ Biologists have used this and similar experiments

century. For the first 40 years of the experiment, Beal checked Source: F. W. Telewski and J. Zeevaart. 2002. Am J Bot 89: 1285–1288.
seed viability every 5 years. H. T. Darlington took over the
experiment in 1915 and extended the sampling period to to estimate the maximum viability of seeds of many species of
10-year intervals beginning in 1920. R. S. Bandurski took over plants under near-natural environmental conditions. Explore how
the experiment when Darlington retired, and extended the these data can be used to make predictions about seed viability in
sampling period to 20 years in 1980, a century after the the Work with the Data exercise.

QUESTION▸ How long do seeds of common plants

experiment began.

METHOD▸
remain viable?

1. Collect 1,000 seeds of each of 23 species of plants.

2. Divide the seeds into 20 lots of 50 seeds of each of the
23 species.
3. Mix each of the lots of seeds with sand, and place the mixtures
in uncapped bottles.
4. Bury the bottles upside down (to prevent the entry of water) on
a marked, sandy knoll.

M. H. Siddall
5. At regular intervals, excavate a bottle and check the viability of
its contents.

Work with the Data

The data presented in this experiment can be used to make KEY CONCEPT
predictions about seed viability. Use the data to answer the 27.1 Recap and Assess
following questions.
1. Calculate the percent of viable seeds for these three Today’s living seed plants fall into two major groups, the gymno-
sperms and the angiosperms. Pollen grains, seeds, and wood are
species in years 50–100 and graph seed survivorship
major evolutionary innovations of the seed plants. Protection of
as a function of time buried.
embryos is a hallmark of seed plants.
2. No seeds of the first two species were viable after 90 1. Explain the importance of pollen in freeing seed plants from
years of the experiment. Assume 100% seed viability dependence on liquid water.
at the start of the experiment (year 0), and predict from
2. How do seeds and seed dormancy help plants survive in highly
your graph the approximate year when you think the last
seasonal environments?
Verbascum blattaria seeds will germinate.
3. What part did the evolution of wood play in changing the
3. What factors do you think might influence the differences terrestrial landscape?
among the species in long-term seed viability?

Go to Achieve for a companion The seed ferns have long been extinct, but the surviving seed
Data in Depth exercise. plants have been remarkable successes. After the seed ferns, the
gymnosperms were the next group of plants to dominate terres-
trial environments.

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 KEY CONCEPT 27.2 Once Dominant Gymnosperms Still Thrive in Some Environments

during the Cenozoic (see Figure 27.1). Although they now number
KEY CONCEPT Once Dominant fewer than 1,200 living species, gymnosperms are second only to
Gymnosperms Still Thrive
27.2 in Some Environments
the angiosperms in their dominance of the terrestrial environment.

There are four major groups of living gymnosperms

Learning Objectives The living gymnosperms can be divided into four major groups:
27.2.1 Describe and contrast the roles of male and female 1. Cycads are palmlike plants of the tropics and subtropics
gametophytes in reproduction. (Figure 27.6A). Of the present-day gymnosperms, the cycads
27.2.2 Summarize the functions of cones in conifers. are probably the earliest-diverging clade. There are about 300
27.2.3 Describe the role of fire in the life history of the species, some of which grow as tall as 20 meters. The tissues
lodgepole pine. of many species are highly toxic to humans if ingested.
2. Ginkgos, common during the Mesozoic, are represented
today by a single species, Ginkgo biloba, the maidenhair tree
The gymnosperms are seed plants Cycads
(Figure 27.6B). There are both male (microsporangiate) and
that do not form flowers or fruits. Ginkgos
female (megasporangiate) maidenhair trees. The difference
Gy m nosperms (wh ich mea ns
Gnetophytes is determined by X and Y sex chromosomes, as in humans.
“naked-seeded”) are so named
Few other plants have distinct sex chromosomes.
because their ovules and seeds, Conifers
unlike those of angiosperms, are not 3. Gnetophytes number about 90 species in three very different
Angiosperms
protected by ovary or fruit tissue. genera, which share certain characteristics analogous to
Gymnosperms dominated the terrestrial world through the Meso- ones found in the angiosperms. One of the gnetophytes is
zoic but were replaced across large areas of Earth by flowering plants Welwitschia mirabilis (Figure 27.6C), a long-lived desert

Cycads

(A) Encephalartos sp. Ginkgos (B) Ginkgo biloba

Gnetophytes

Conifers

Angiosperms
Courtesy of Andrew D. Sinauer

David McIntyre
(C) Welwitschia mirabilis (D) Pinus longaeva © Richard G Smith/Shutterstock.com
© iStock.com/namibelephant

Figure 27.6 Diversity among the Gymnosperms (A) Many cycads straplike leaves of Welwitschia mirabilis, a gnetophyte, grow throughout
have growth forms that resemble those of both ferns and palms, the life of the plant, breaking and splitting as they grow. (D) Conifers
although cycads are not closely related to either group. (B) The char- dominate many types of landscapes in the Northern Hemisphere. Bristle-
acteristic broad leaves of the maidenhair tree. (C) The two permanent cone pines such as these are the longest-lived individual trees known.

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 CHAPTER 27 The Evolution of Seed Plants

plant with just two permanent straplike leaves, which split

of the world. All these trees belong to one group of gymnosperms:
into many pieces that sprawl across the sand.
the conifers, or cone-bearers.
4. Conifers are by far the most abundant of the gymnosperms. Male and female cones contain the reproductive structures of
There are about 700 species of these cone-bearing plants, conifers. The female (seed-bearing) cone is known as a megastro-
including the pines, spruces, and redwoods (Figure 27.6D). bilus (plural megastrobili). An example of a familiar megastrobilus
With the exception of the gnetophytes, the living gymnosperm is the woody cone of pine trees. The seeds in a megastrobilus are
groups have only tracheids as water-conducting and support cells protected by a tight cluster of woody scales, which are modifica-
in the xylem. Most gymnosperms lack the vessel elements and fibers tions of branches extending from a central axis (Figure 27.7A). The
(cells specialized for water conduction and support, respectively) typically much smaller male (pollen-bearing) cone is known as a
that are found in angiosperms. While the gymnosperm water-trans- microstrobilus. The microstrobilus is typically herbaceous rather
port and support system may thus seem somewhat less efficient than woody, as its scales are composed of modified leaves, beneath
than that of the angiosperms, it serves some of the largest trees which are the pollen-bearing microsporangia (Figure 27.7B).
known. The coastal redwoods of California are the tallest gymno- The life cycle of a pine illustrates reproduction in gymnosperms
sperms, with some individuals growing to well over 100 meters tall. (Figure 27.8). As in other seed plants, conifers have male gameto-
During the Permian, as environments became warmer and dryer, phytes in the form of pollen grains, which frees the plant completely
the conifers and cycads flourished. Gymnosperm forests changed from its dependence on liquid water for fertilization. Wind, rather
over time as the gymnosperm groups evolved. Gymnosperms domi- than water, assists conifer pollen grains in their first stage of travel
nated the Mesozoic, during which the continents drifted apart and from the microstrobilus to the female gametophyte inside a cone. A
large dinosaurs lived. Gymnosperms were the principal trees in all pollen tube provides the sperm with the means for the last stage of
forests until about 65 million years ago, and even today conifers travel by elongating through maternal sporophytic tissue. When the
are the dominant trees in many forests, especially at high latitudes pollen tube reaches the female gametophyte, it releases two sperm,
and elevations. The oldest living single organism on Earth today is one of which degenerates after the other unites with an egg. Union
a gymnosperm in California—a bristlecone pine (Pinus longaeva) of sperm and egg results in a zygote. Mitotic divisions and further
that germinated about 5,000 years ago, at about the time the ancient development of the zygote result in an embryo.
Egyptians were starting to develop writing. The megasporangium, in which the female gametophyte will
form, is enclosed in a layer of sporophytic tissue—the integu-
Conifers have cones and lack swimming sperm ment—that will eventually develop into the seed coat that protects
The great Douglas fir and cedar forests found in the northwestern the embryo. The integument, the megasporangium inside it, and
United States and the massive boreal forests of pine, fir, and spruce the tissue attaching it to the maternal sporophyte constitute the
of northern Eurasia and North America, as well as on the upper ovule. The pollen grain enters through a small opening in the
slopes of mountain ranges elsewhere, rank among the great forests integument at the tip of the ovule, the micropyle.

(A) Female cones (megastrobili) in Pinus contorta (B) Male cones (microstrobili) in Pinus contorta

Cross section of Cross section of

a megastrobilus a microstrobilus
Herbaceous
scale
Woody Pollen-
scale containing
microspores
Seed
Central
Central axis
axis

© Scenics & Science/Alamy Stock Photo

© Scenics & Science/Alamy Stock Photo

Male cones,
Female cones, or microstrobili
or megastrobili © Gunter Marx/Alamy Stock Photo

© Fritz Pölking/Blickwinkel/AGE Fotostock

Figure 27.7 Female and Male Cones (A) The woody scales of female cones (megastrobili) are
modified branches. (B) The herbaceous scales of male cones (microstrobili) are modified leaves.

27_Life12e_Ch 27.indd 8 12/4/19 9:42 AM

 KEY CONCEPT 27.2 Once Dominant Gymnosperms Still Thrive in Some Environments

Scale of
The sporophyte is The same plant has both megastrobilus
an enormous tree. pollen-producing microstrobili
and egg-producing megastrobili.
Integument

Immature Megasporocyte
megastrobilus
Meiosis
Functional
Ovule megaspore

Megasporangium

Pollen
chamber
Microstrobili

Meiosis

Microsporangium

Scale of Micropyle
microstrobilus
Microspores

Sporophyte Pollen
(10–100 m) grain

Haploid (n)
Seed coat Gametophyte
generation
Female gametophyte
(provides nutrition for Diploid (2n) Female
developing embryo) Sporophyte gametophyte
generation
Embryo
Egg

Sperm

Winged Male
gametophyte
seed
(pollen tube)
Seed
Mature coat
megastrobilus Fertilization
Scale of Zygote
megastrobilus
The gametophyte
is tiny compared
Wing
with the sporophyte.

Seed

Figure 27.8 The Life Cycle of a Pine Tree In conifers and other View in Achieve
gymnosperms, the gametophytes are small and nutritionally dependent Animation 27.1 Life Cycle of a Conifer
on the sporophyte generation. View in Achieve
Activity 27.2 Life Cycle of a Conifer

Most conifer ovules (which will develop into seeds after fertil- within the cone. Some pines, such as the lodgepole pine (Pinus
ization) are borne exposed on the upper surfaces of the scales of contorta), have such tightly closed cones that only fire suffices to
the cone (megastrobilus). The only protection of the ovules comes split them open and release the seeds. These species are said to
from the scales, which are tightly pressed against one another be fire-adapted, and fire is essential to their reproduction. A fire

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 CHAPTER 27 The Evolution of Seed Plants

KEY CONCEPT Flowers and Fruits Led to

Increased Diversification
27.3 of Angiosperms
Learning Objectives Cycads

27.3.1 Distinguish between the two different roles of sperm

Ginkgos
in angiosperm double fertilization.

© robert cicchetti/Alamy Stock Photo

Gnetophytes
27.3.2 Explain the difference between pollination and
fertilization. Conifers
27.3.3 Predict the likely modes of seed dispersal based on
Angiosperms
the type of fruit a plant produces.
27.3.4 Outline the functions of flowers, fruits, and seeds.

Figure 27.9 From Devastation, New Life A stand of lodgepole The most obvious feature de- Cycads
pines in Yellowstone National Park. The mature trees were destroyed fining the angiosperms is the Ginkgos
by a forest fire in 1988. However, the fire released large numbers of flower, which is their sexual
seeds from cones, and now many young lodgepole pine trees are grow- Gnetophytes
structure. Production of fruits
ing in the burn area.
is also a synapomorphy (shared Conifers
derived trait) of angiosperms.
Angiosperms
devastated lodgepole pine forests in Yellowstone National Park in After fertilization, the ovary of
1988, but also released large numbers of seeds from cones. As a a flower (together with the seeds
result, large numbers of lodgepole pine seedlings are now emerg- it contains) develops into a fruit that protects the seeds and can
ing in the burn area (Figure 27.9). promote seed dispersal. As we will see, both flowers and fruits
About half of all conifer species have soft, fleshy modifications afford major reproductive advantages to angiosperms. These ad-
of the cones that envelop their seeds. Some of these modified tis- vantages led to the dominance of the terrestrial environment by
sues are fleshy, fruitlike cones, as in junipers. Others are fruitlike angiosperms in the Cenozoic (see Figure 27.1).
extensions of the seeds, called arils, as in yews. These tissues, al-
though often mistaken for “berries,” are not true fruits. As you will Angiosperms have many shared derived traits
see in Key Concept 27.3, true fruits are the plant’s ripened ovaries, The name angiosperm (“enclosed seed”) is derived from another
which are absent in gymnosperms. Nonetheless, the fleshy tissues distinctive trait of flowering plants that is related to the forma-
that surround many conifer seeds serve a similar purpose as that tion of fruits: the ovules and seeds are enclosed in a modified
of the fruits of flowering plants, acting as an enticement for seed- leaf called a carpel. Besides protecting the ovules and seeds,
dispersing animals. Animals eat these fleshy tissues and disperse the carpel often interacts with incoming pollen to prevent self-
the seeds in their feces, often depositing the seeds considerable fertilization, thus favoring cross-fertilization and increasing
distances away from the parent plant. genetic diversity.
The female gametophyte of the angiosperms is even more
KEY CONCEPT
reduced than that of the gymnosperms, usually consisting of only
27.2 Recap and Assess seven cells (see Figure 27.16). Thus the angiosperms represent
Living gymnosperms can be divided into four major groups: the current extreme of the trend we have traced throughout the
cycads, ginkgos, gnetophytes, and conifers. Although they have evolution of the vascular plants: the sporophyte generation be-
declined since the Mesozoic, gymnosperms are still the dominant comes larger and more independent of the gametophyte, while
trees in some areas, especially at high elevations and latitudes. the gametophyte generation becomes smaller and more depen-
All gymnosperms are woody and have seeds that are not protect-
ed by ovaries, although some have fleshy, fruitlike cones or exten- dent on the sporophyte.
sions of seeds that entice animals to disperse the seeds. The xylem of most angiosperms is distinguished by the pres-
1. Why is fire necessary for the survival of some gymnosperms? ence of specialized water-transporting cells called vessel elements.
2. Distinguish between the roles of the female gametophyte and These cells are larger in diameter than tracheids and connect with
the pollen grain in the life cycle of a conifer. one another without obstruction, allowing easy water movement. A
3. What is the function of the fleshy cones that surround the second distinctive cell type in angiosperm xylem is the fiber, which
seeds of many gymnosperms? plays an important role in supporting the plant body. Angiosperm
phloem possesses another unique cell type, called a companion
Although gymnosperms still dominate the terrestrial landscape cell. Like the gymnosperms, woody angiosperms show secondary
in some environments, angiosperms (the flowering plants) have growth, increasing in diameter by producing secondary xylem and
become the dominant land plants across much of Earth. secondary phloem.

Life 12e
27_Life12e_Ch 27.indd 10 Oxford University Press 12/4/19 9:42 AM
Dragonfly Media Group
Life12e_UN 27.01.ai Date 07-10-19
 KEY CONCEPT 27.3 Flowers and Fruits Led to Increased Diversification of Angiosperms

A more comprehensive list of angiosperm synapomorphies, integuments), is called the ovary. The stalk at the top of the
then, includes the following (some of these traits will be discussed carpel is the style, and the terminal surface that receives pollen
later in this chapter): grains is the stigma. Two or more fused carpels, or a single carpel
• Flowers if only one is present, are also called a pistil.
• Fruits
In addition, many flowers have specialized sterile (non-spore-
bearing) leaves. The inner ones are called petals (collectively,
• Ovules and seeds enclosed in a carpel the corolla) and the outer ones sepals (collectively, the calyx).
• Highly reduced gametophytes The corolla and calyx can be quite showy and often play roles
• Germination of pollen on a stigma in attracting animal pollinators to the flower. The calyx more
• Double fertilization
commonly protects the immature flower in bud. From base to
apex, these floral organs—sepals, petals, stamens, and carpels—
• Endosperm (nutritive tissue for the embryo) are usually positioned in circular arrangements or whorls and
• Phloem with companion cells attached to a central stalk.
The majority of these traits bear directly on angiosperm repro- The generalized flower in Figure 27.4B has both functional
duction, which is a large factor in the success of this dominant megasporangia and functional microsporangia. Such flowers are
plant group. referred to as perfect (or hermaphroditic). Many angiosperms pro-
duce two types of flowers, one with only megasporangia and the
The sexual structures of angiosperms are flowers other with only microsporangia. Consequently, either the stamens
Flowers come in an astonishing variety of forms—just think of or the carpels are nonfunctional or absent in a given flower, and
some of the flowers you recognize. Flowers may be single, or they the flower is referred to as imperfect.
may be grouped together to form an inflorescence. Different fami- Species such as corn or birch, in which both megasporangiate
lies of flowering plants have characteristic types of inflorescences, (female) and microsporangiate (male) flowers occur on the same
such as the compound umbels of the carrot family (Figure 27.10A), plant, are said to be monoecious (“one-housed”—but, it must be
the heads of the aster family (Figure 27.10B), and the spikes of added, one house with separate rooms). Complete separation of
many grasses (Figure 27.10C). imperfect flowers occurs in some other angiosperm species, such
If you examine any familiar flower, you will notice that the as willows and date palms; in these species, an individual plant
outer parts look somewhat like leaves. In fact, all the parts of a produces either flowers with stamens or flowers with carpels, but
flower are modified leaves. The diagram in Figure 27.4B rep- never both. Such species are said to be dioecious (“two-housed”).
resents a generalized flower (for which there is no exact coun-
terpart in nature). The structures bearing microsporangia are Flower structure has evolved over time
called stamens. Each stamen is composed of a filament bearing The flowers of the earliest-diverging clades of angiosperms have
an anther that contains the pollen-producing microsporangia. a large and variable number of tepals (undifferentiated sepals and
The structures bearing megasporangia are called carpels. The petals), carpels, and stamens (Figure 27.11A). Evolutionary change
swollen base of the carpel, containing one or more ovules (each within the angiosperms has included some striking modifications
containing a megasporangium surrounded by two protective of this early condition: reductions in the number of each type of

(A) Heracleum mantegazzianum (B) Zinnia elegans (C) Sorghum halepense

Flowers
© John N. A. Lott/Biological Photo Service
© Colin Underhill/Alamy Stock Photo

© Nigel Cattlin/Alamy Stock Photo

Umbel

Compound umbel

Ray flowers Disc flowers Anthers of spikelet Stigma of spikelet

Figure 27.10 Inflorescences (A) The inflorescence of the giant flower; the central portion of the head consists of dozens to hundreds
hogweed, a member of the carrot family, is a compound umbel. Each of disc flowers. (C) Some grasses, such as Johnson grass, have inflo-
umbel bears flowers on stalks that arise from a common center. rescences called spikes, which are composed of many smaller groups
(B) Zinnias are members of the aster family; their inflorescence is a of flowers, or spikelets.
head. Within the head, each of the long, petal-like structures is a ray

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 CHAPTER 27 The Evolution of Seed Plants

© imageBROKER/Alamy Stock Photo

David McIntyre
(A) Nymphaea sp. (B) Paphiopedilum sp.

Figure 27.11 Flower Form and Evolution (A) A water lily numerous, and attached at their bases. (B) Orchids such as this
shows the major features of early flowers: it is radially symmetri- Venus slipper have a bilaterally symmetrical structure that evolved
cal, and the individual tepals, stamens, and carpels are separate, much later than radial flower symmetry.

floral organ to a fixed number, differentiation of petals from se-

(A) Carpel evolution pals, and changes in symmetry from radial (as in a lily or magnolia)
to bilateral (as in a sweet pea or orchid), often accompanied by an
1 According to one 2 In the course of 3 At the end of the
extensive fusion of parts (Figure 27.11B).
theory, the carpel evolution, leaf edges sequence, three
began as a curled inward and carpels have fused According to one hypothesis, the first carpels to evolve were
modified leaf finally fused. to form a three- leaves with marginal sporangia, folded but incompletely closed.
bearing ovules. chambered ovary. Early in angiosperm evolution, the carpels fused with one an-
other, forming a single, multichambered ovary (Figure 27.12A).
Ovules Fused In some flowers, the other floral organs are attached at the top of
carpel
the ovary rather than at the bottom as in Figure 27.4B. The sta-
mens of the most ancient flowers may have been leaflike (Figure
27.12B), with little resemblance to the stamens of the general-
Modified
ized flower seen in Figure 27.4B.
leaflike Why do so many flowers have carpels with long styles and an-
structure thers with long filaments? Natural selection has favored length
Cross in both of these floral organs, probably because length increases
sections the likelihood of successful pollination. Long filaments may bring
the anthers into contact with insect bodies, or they may place the
(B) Stamen evolution anthers in a better position to catch the wind. Similar arguments
apply to long styles.
1 The leaflike portion of the structure 2 …until only the micro-
was progressively reduced… sporangia remained. A perfect flower represents a compromise of sorts. On the one
hand, by attracting a pollinating bird or insect, the plant is attend-
Austrobaileya sp. Magnolia Lily ing to both its female and male functions with a single flower type,
Modified whereas plants with imperfect flowers must create that attraction
leaf twice—once for each type of flower. On the other hand, the perfect
flower can favor self-pollination, which is usually disadvantageous.
Anther
Another potential problem is that the female and male functions
might interfere with each other—for example, the stigma might be
Sporangia Filament so placed as to make it difficult for pollinators to reach the anthers,
Cross thus reducing the export of pollen to other flowers.
sections Might there be a way around these problems? One solution is
seen in the bush monkeyflower (Mimulus aurantiacus), which
Figure 27.12 Carpels and Stamens Evolved from Leaflike
is pollinated by hummingbirds. Its flower has a stigma that ini-
Structures (A) Possible stages in the evolution of a carpel from a
more leaflike structure. (B) The stamens of three modern plants show tially serves as a screen, hiding the anthers (Figure 27.13). Once
three possible stages in the evolution of that organ. (It is not implied a hummingbird touches the stigma, however, one of the stigma’s
that these species evolved from one another; their structures simply two lobes is retracted, so that subsequent hummingbird visitors
illustrate the possible stages.) pick up pollen from the previously screened anthers. Thus the first

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 KEY CONCEPT 27.3 Flowers and Fruits Led to Increased Diversification of Angiosperms

Stigma open Stigma

(access to
Experiment
Anthers
anthers blocked) Figure 27.14 The Effect of Stigma Retraction in
Monkeyflowers
Original Paper: A. E. Fetscher. 2001. Resolution of male-female
conflict in an hermaphroditic flower. Proc R Soc Lond B Biol Sci
Filaments Style 268: 525–527.
Elizabeth Fetscher’s experiments showed that the unusual
stigma retraction response to pollination in monkeyflowers
(illustrated in Figure 27.13) enhances the dispersal of pollen
Stigma retracted

HYPOTHESIS▸ The stigma-retraction response in Mimulus

(anthers to other monkeyflowers.
accessible)
Access aurantiacus increases the likelihood that an individual flower’s
route pollen will be exported to another flower once pollen from

METHOD▸
another flower has been deposited on its stigma.

1. Set up three groups of monkeyflower arrays. Each array

consists of one pollen-donor flower and multiple pollen-
recipient flowers (with the anthers removed to prevent
pollen donation).
Figure 27.13 An Unusual Way to Prevent Selfing Both long
stamens and long styles facilitate cross-pollination, but if these male 2. In control arrays, the stigma of the pollen donor is allowed
and female structures are too close to each other, the likelihood of to function normally.
(disadvantageous) self-pollination increases. In Mimulus aurantiacus, 3. In one group of experimental arrays, the stigma of the
the stigma is initially open, blocking access to the anthers. A humming- pollen donor is permanently propped open (blocking
bird’s touch as it deposits pollen on the stigma causes one lobe of the access to the anthers).
stigma to retract, creating a path to the anthers and allowing pollen 4. In a second group of experimental arrays, the stigma of
dispersal by subsequent hummingbird visitors. (After A. E. Fetscher. the pollen donor is artificially sealed closed (allowing
2001. Proc R Soc Lond B Biol Sci 268: 525–529.) access to the anthers).
5. Allow hummingbirds to visit the arrays, then count the
pollen grains transferred from each donor flower to the

RESULTS▸ Error bars indicate standard error of the mean; the

bird to visit the flower transfers pollen from another plant to the recipient flowers in the same array.
stigma, eventually leading to fertilization. Later visitors pick up
pollen from the now-accessible anthers, fulfilling the flower’s male three groups are significantly different at P<0.05.
function. Figure 27.14 describes the experiment that revealed the
function of this mechanism. Almost twice as much pollen Experimentally sealing
was exported from control stigmas in the closed
Angiosperms have coevolved with animals flowers as from flowers with position resulted in
stigmas propped open. greater pollen dispersal.
Whereas most gymnosperms are pollinated by wind, most angio-
sperms are pollinated by animals. The many different mutualistic
pollination relationships between plants and animals are vital to 350
Number of pollen grains deposited

both parties. 300

Connect the Concepts Mutualisms benefit both partners in a
250
relationship, as described in Chapter 54.
on recipients

Many flowers entice animals to visit them by providing food 200

rewards. Pollen grains themselves sometimes serve as food for
150
animals. In addition, some flowers produce a sugary fluid called
nectar as a pollinator attractant, and some of these flowers have 100
specialized structures to store and distribute it. In the process of
visiting flowers to obtain nectar or pollen, animals often carry 50
pollen from one flower to another or from one plant to another. 0
Controls Stigmas Stigmas
Thus, in their quest for food, the animals contribute to the ge- propped open sealed closed

CONCLUSION▸ The stigma-retraction response enhances the

netic diversity of the plant population. Insects, especially bees, are
among the most important pollinators. Other major pollinators
include some species of birds and bats. male function of the flower (dispersal of pollen) once the female
Media Clip 27.2 Pollen Transfer by a Bat function (receipt of pollen) has been performed.
Life12e.com/mc27.2

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 CHAPTER 27 The Evolution of Seed Plants

For more than 150 million years, angiosperms and their ani- mammoths, osage orange trees survived in large part because
mal pollinators have coevolved in the terrestrial environment. early humans used the wood of this species for making bows for
The animals have affected the evolution of the plants, and the hunting, dispersing the fruits to new areas in the process. Some
plants have affected the evolution of the animals. Flower struc- other species that depended on now-extinct mammals to disperse
ture has become incredibly diverse under these selection pres- their seeds, however, may be slowly declining toward extinction.
sures. Some of the products of coevolution are highly specific.
For example, the flowers of some yucca species are pollinated by The angiosperm life cycle produces diploid zygotes
only one species of yucca moth, and that moth may exclusively nourished by triploid endosperms
pollinate just one species of yucca. Such specific relationships Like all seed plants, angiosperms are heterosporous. As you have
provide plants with a reliable mechanism for transferring pollen seen, their ovules are contained within carpels rather than being
only to members of their own species. exposed on the surfaces of scales, as in most gymnosperms. The
Most plant–pollinator interactions are much less specific. In male gametophytes, as in the gymnosperms, are pollen grains.
most cases, many different animal species pollinate the same plant Pollination in the angiosperms consists of the arrival of a male
species, and the same animal species pollinates many different gametophyte—a pollen grain—on a receptive surface in a flower
plant species. However, even these less specific interactions have (the stigma). As in the gymnosperms, pollination is the first in a
developed some specialization. Bird-pollinated flowers are often series of events that results in the formation of a seed. The next
red and odorless. Many insect-pollinated flowers have character- event is the growth of a pollen tube extending to the female game-
istic odors. Bee-pollinated flowers may have conspicuous mark- tophyte. The third event is a fertilization process that, in detail, is
ings, called nectar guides, that are conspicuous only to animals, unique to the angiosperms (Figure 27.16).
such as bees, that can see colors in the ultraviolet region of the In nearly all angiosperms, two male gametes, contained in a
spectrum (Figure 27.15). single male gametophyte, participate in fertilization. The nucle-
The fruits of some plants that are still around today originally us of one sperm combines with that of the egg to produce a dip-
evolved to attract large frugivorous animals—many of which went loid zygote, the first cell of the sporophyte generation. In most
extinct in the Pleistocene. For example, the large round fruits of angiosperms, the other sperm nucleus combines with two other
the osage orange tree attracted wooly mammoths, which ate haploid nuclei of the female gametophyte to form a cell with a
the fruits and dispersed the seeds. After the extinction of the triploid (3n) nucleus. That cell, in turn, gives rise to triploid
tissue, the endosperm, which nourishes the embryonic sporo-
phyte during its early development. This process, in which two
Argentina anserina
fertilization events take place, is known as double fertilization.
In some angiosperms, additional haploid nuclei are incorporated
to form even higher ploidy levels in the endosperm, or the second
sperm fuses with only one haploid nucleus, resulting in diploid
endosperm.
As Figure 27.16 shows, the zygote develops into an embryo,
which consists of an embryonic axis (the “backbone” that will
become a stem and a root) and one or two cotyledons, or “seed
leaves.” The cotyledons have different fates in different plants. In
many, they serve as absorptive organs that take up and digest the
endosperm. In others, they enlarge and become photosynthetic
when the seed germinates. Often they play both roles.
The ovule develops into a seed containing the products of the
double fertilization that characterizes angiosperms: a diploid zy-
gote and a triploid endosperm (see Figure 27.16). The endosperm
Both photos © Bjorn Rorslett/Science Source

serves as storage tissue for starch or lipids, proteins, and other

substances that will be needed by the developing embryo.

Fruits aid angiosperm seed dispersal

Fruits typically aid in seed dispersal. Fruits may attach to or be
eaten by an animal. The animal is then likely to move, after which
the seeds may fall off or be defecated. Fruits are not necessar-
ily fleshy. Fruits can also be hard and woody, or small and have
Figure 27.15 See Like a Bee To normal human vision, the modified structures that allow the seeds to be dispersed by wind
petals of silverweed appear solid yellow. Ultraviolet photography or water (Figure 27.17).
reveals patterns that attract bees to the central region, where A fruit may consist of only the mature ovary and its seeds, or
pollen and nectar are located. it may include other parts of the flower or structures associated

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 KEY CONCEPT 27.3 Flowers and Fruits Led to Increased Diversification of Angiosperms

Flower of mature sporophyte

Anther

Seedling

Ovary
Seed
Cotyledons Ovule
Double fertilization Megasporocyte (2n)
results in a 2n zygote
and 3n endosperm. Megasporangium
Endosperm
Embryo

Endosperm Anther
nucleus (3n)

Microsporocyte
Zygote (2n)

Diploid (2n)
Haploid (n)
Double Fertilization Meiosis

Pollen grain
(male gametophyte, n)
Microspores (4)

Pollen grain

Pollen tube

Three of the four

Surviving
megaspores that
megaspore (n)
result from meiosis
degenerate.

Sperm Polar nuclei (2)

The typical angiosperm

female gametophyte (n)
consists of seven cells.
Egg

Figure 27.16 The Life Cycle of an Angiosperm Double fertiliza- View in Achieve
tion results in triploid endosperm in most species of angiosperms. One Animation 27.2 Life Cycle of an Angiosperm
sperm nucleus fertilizes the egg to form the zygote, while the other
combines with the two polar nuclei to form the endosperm.

with it. A simple fruit is one that develops from a single carpel inflorescence). Fruits derived from parts in addition to the carpel
or several fused carpels, such as a plum or cherry. A raspberry is and seeds are called accessory fruits—examples are apples, pears,
an example of an aggregate fruit—one that develops from sev- and strawberries.
eral separate carpels of a single flower. Pineapples and figs are Media Clip 27.3 Flower and Fruit Formation
examples of multiple fruits, formed from a cluster of flowers (an Life12e.com/mc27.3

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 CHAPTER 27 The Evolution of Seed Plants

(A) (B) (C)

Brian Jackson/Alamy Stock Photo

© Masayuki/Shutterstock.com
© iStock.com/Leonsbox

(D) (E) (F)

© aniad/Shutterstock.com

© iStock.com/Dejan Kolar
© iStock.com/mazzzur

Figure 27.17 Fruits Come in Many Forms (A) The single seeds in- Q: The large incisors of rodents help these animals penetrate
side the simple fruits of Bing cherries are dispersed by animals. (B) Each the shells of woody nuts, which they consume as food. But
seed of the horse chestnut is covered by a hard, woody fruit that allows unlike many animals that eat fruits without digesting the
it to survive drought. Although such fruits are commonly called “nuts,” seeds, rodents destroy the seeds when they eat them.
this is a culinary rather than a biological term. (C) The highly reduced So how do rodents aid in dispersing the seeds of
simple fruits of dandelions are dispersed by wind. (D) A multiple fruit, the nut-bearing plants?
pineapple (Ananas comosus), has become one of the most economically
significant fruit crops of the tropics. (E) An aggregate fruit (blackberry).
(F) An accessory fruit (pear).

Amborella

Common
ancestor of
angiosperms Water lilies

Carpels; endosperm;
seeds in fruit; reduced Star anise
gametophytes; double
fertilization; flowers;
phloem with companion
cells Magnoliids
Transitional Vessel Single
tracheid elements cotyledon
vessel
elements Carpel margins Monocots
fused by tissue
Figure 27.18 Evolutionary Relationships among the connection
Sepals and petals
Life 12e Angiosperms Recent analyses of many angiosperm genes of two whorls Eudicots
Oxford University Press
have clarified the relationships among the major groups. Pollen with
Dragonfly Media Group
Life12e_27.17.ai Date 07-10-19 three grooves

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 KEY CONCEPT 27.3 Flowers and Fruits Led to Increased Diversification of Angiosperms

(A) Amborella trichopoda (B) Nymphaea sp. (C) Illicium floridanum

© Rob & Ann Simpson/Visuals Unlimited, Inc.

courtesy of the U. Massachusetts

© Yuriy Kulik/Shutterstock.com
Biology Department Greenhouses

David McIntyre,

© foto76/Shutterstock.com
Sterile stamens

© iStock.com/Boogich

(D) Magnolia sp. (E) Aristolochia ringens

Figure 27.19 Monocots and Eudicots Are Not the Only Surviving was the next to diverge after Amborella. (C) Star anise and its relatives
Angiosperms (A) Amborella, a shrub, is sister to the remaining extant belong to another early-diverging angiosperm clade. (D, E) The largest
angiosperms. Notice the sterile stamens on this female flower, which may clade other than the monocots and eudicots is the magnoliid complex,
serve to lure insects that are searching for pollen. (B) The water lily clade which includes magnolias and the group known as “Dutchman’s pipe.”

Recent analyses have revealed the phylogenetic star anise and its relatives (Figure 27.19C), and the magnoliids
relationships of angiosperms (Figure 27.19D,E). The magnoliids include many familiar and useful
Figure 27.18 shows the relationships among the major angiosperm plants, such as avocados, cinnamon, black pepper, and magnolias.
clades. Recent molecular and morphological analyses have sup- The two largest clades—the monocots and the eudicots—in-
ported the hypothesis that the sister group of remaining flowering clude the great majority of angiosperm species. The monocots are
plants is a single species of the genus Amborella (Figure 27.19A). so called because they have a single embryonic cotyledon, whereas
This woody shrub with cream-colored flowers lives only on New the eudicots have two.
Caledonia, an island in the South Pacific. Other early-branch- Representatives of the two largest angiosperm clades are every-
ing angiosperm groups include the water lilies (Figure 27.19B), where. The monocots (Figure 27.20) include grasses, cattails, lilies,

(A) Tulipa sp. and Narcissus sp. (B) Oryza sativa

© Craig Roberts/Alamy Stock Photo

© Prisma Bildagentur AG/

Alamy Stock Photo

© blickwinkel/Alamy Stock Photo

© Images & Stories/
Alamy Stock Photo

(C) Posidonia oceanica (D) Phoenix dactylifera

Figure
Life 12e 27.20 Monocots (A) Monocots include many popular garden Neptune’s grass form “meadows” in the shallow, sunlit waters of the
flowers University
Oxford such as these
Presstulips (pink and white) and daffodils (yellow). world’s oceans. (D) Palms are among the few monocot trees. Date palms
Dragonfly Media
(B) Monocot Group
grasses such as rice feed the world; wheat, sugarcane, like these are a major food source in some areas of the world.
Life12e_27.19.ai Date
and corn (maize) are also 07-15-19
grasses. (C) Seagrasses such as this

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 CHAPTER 27 The Evolution of Seed Plants

(A) Malus sp. orchids, and palms. The eudicots (Figure 27.21) include the vast
majority of familiar seed plants, including most herbs (i.e., nonwoody

© John N. A. Lott/Biological Photo Service

plants), vines, trees, and shrubs. Among the eudicots are such diverse
plants as oaks, willows, beans, snapdragons, roses, and sunflowers.
KEY CONCEPT
27.3 Recap and Assess
Angiosperms are the dominant terrestrial plants of the Cenozoic.
The synapomorphies of angiosperms include flowers, fruit, car-
pels, double fertilization, and endosperm. Most angiosperms also
possess distinctive cells in the xylem and phloem. The largest
angiosperm clades are the monocots and the eudicots.
1. What are the respective roles of the two sperm in double
fertilization in angiosperms?
(B) Banksia 2. Explain the difference between pollination and fertilization.
coccinea 3. What are some of the differences between fruits that are
dispersed by attracting animals to a food resource and those
that are dispersed by attaching themselves to animals? Give
examples of each.
4. What are the different functions of flowers, fruits, and seeds?
5. Many fleshy fruits attract animals, which eat the fruit and then
(C) Escobaria vivipara disperse the seeds in their feces. If all other factors are equal,
large seeds have a better chance of producing a successful
seedling than small seeds. So why isn’t there selection for
larger seeds in the fruits of all plants? In one study in Peru,
the feces of the spider monkey Ateles paniscus were found to
contain seeds from 71 species of plants. After eating fruit,
the monkeys usually travel some distance before defecating,
thus dispersing any undigested seeds. If monkey feces are
Courtesy of David Hillis

left undisturbed on the forest floor, rodents eat and destroy

the vast majority of the seeds in the feces. To germinate
Cygnis insignis

successfully, the seeds in spider monkey feces need to

be buried by dung beetles, which makes the discovery and
destruction of seeds by rodents much less likely.
(D) Rafflesia arnoldii Ellen Andresen hypothesized that dung beetles were more
likely to remove larger than smaller seeds from spider monkey
dung before burying the dung. She added plastic beads of
various diameters to spider monkey dung (to simulate seeds)
and measured the percentage of beads buried with the dung
by the beetles. Use her data to answer the questions below.

Bead diameter (mm) 2 4 6 8 10 12

Percent buried 100 76 52 39 20 4
Source: E. Andresen. 1999. Biotropica 31: 145–158.
© A & J Visage/Alamy Stock Photo

a. Plot bead size (the independent variable) versus percent of

beads buried by dung beetles (the dependent variable).
b. Calculate a regression line for the relationship shown in your
graph (see Appendix B). Approximately what percent of beads
with a diameter of 5 mm would you predict would be buried
by the beetles? What about beads 14 mm in diameter?
c. What other factors besides size might influence the prob-
ability of seed burial by dung beetles? Can you design an
Figure 27.21 Eudicots (A) Eudicots include many trees, such experiment to test your hypotheses?
as this crabapple tree. (B) Scarlet Banksia is a species of an Austra-
d. Describe how changes in the population sizes of spider
lian genus of eudicots that attracts a wide diversity of pollinators by
monkeys, rodents, and dung beetles would affect the
producing large quantities of nectar. (C) Cacti comprise a large group
reproductive success of various plant species.
of eudicots, with about 1,500 species in the Americas. Many, such as
this spinystar cactus, bear large flowers for a brief period of each year.
(D) Rafflesia arnoldii, found in the rainforests of Indonesia, bears the
The remarkable diversity of the seed plants has been shaped by
largest flower in the world. The flower lives as a parasite on tropical both biotic and abiotic components of the environments to which
vines and has lost its leaf, stem, and even root structures. It smells they have adapted. In turn, land plants—and seed plants in par-
like decaying meat, which attracts its fly pollinators. ticular—shape their environments.

Life 12e
Oxford University Press
Dragonfly Media Group
Life12e_27.21.ai Date 07-10-19

27_Life12e_Ch 27.indd 18 12/4/19 9:42 AM

 KEY CONCEPT 27.4 Plants Play Critical Roles in Terrestrial Ecosystems

KEY CONCEPT Plants Play Critical Roles in

27.4 Terrestrial Ecosystems
Learning Objectives
27.4.1 Describe how seed plants have been used for
medicinal purposes.
27.4.2 Summarize the ecological services that angio-

© Calvin Chan/Alamy Stock Photo

sperms perform.
27.4.3 Summarize the main plants that humans use
for food.

Plants make profound contributions to ecosystem services—pro-

cesses by which the environment maintains resources that benefit
human society. Once life moved onto land, it was largely plants
that shaped the terrestrial environment. Figure 27.22 Plants Prevent Erosion When forest vegetation was
Plants produce oxygen and remove carbon dioxide from the cleared on these hillsides in Malaysia, landslides and extensive soil ero-
sion quickly followed. Adjacent forested areas did not have landslides.
atmosphere, and they play important roles in forming soils and
renewing soil fertility. Plant roots help hold soil in place, provid-
ing protection against erosion by wind and water (Figure 27.22). century, Spanish priests in Peru became aware that the native
Plants also moderate the local climate in various ways, such as population used the bark of local Cinchona trees to treat fevers.
by increasing humidity, providing shade, and blocking wind. All The priests successfully used the bark to treat malaria. Word of
of these ecosystem services permit a great diversity of fungi and the medicine spread to Europe, where it was put into use as early
animals to exist on land. as 1631. The active ingredient of Cinchona bark—quinine—was
identified in 1820, and quinine remained the standard malarial
Seed plants have been sources of medicine since remedy well into the twentieth century.
ancient times Since the mid-1900s, many plant-based medicines have been
Although we also use medicines derived from fungi, lichens, and found by systematic testing of plants from all over the world. One
actinobacteria, seed plants are the source of many of our medica-
tions. A few examples of medicines derived from plants are shown
®
example of a medicine discovered in this way is Taxol , an impor-
tant anticancer drug. Among the myriad plant samples that had
in Table 27.1. Even in synthetic pharmaceuticals, the chemical been tested by 1962, extracts of the bark of the Pacific yew (Taxus
structures of the active ingredients are often based on the bio- brevifolia) showed anti-tumor activity in tests against rodent tu-
chemistry of substances isolated from plants. mors. The active ingredient, paclitaxel, was isolated in 1971 and
How are plant-based medicines discovered? Many were dis- tested against human cancers in 1977. After another 16 years,
covered over the millennia by people who lived alongside useful the U.S. Food and Drug Administration approved it for human
plants and discovered the properties of the plants through trial
and error. Some plant-based medicines have come into widespread
®
use, and Taxol is now widely used in treating breast and ovarian
cancers as well as several other types of cancers.
use through the work of ethnobotanists, who study how people use
and view plants in their local environments. This work proceeds Seed plants are our primary food source
all over the globe today. An older example of this approach is the Plants are primary producers. They trap energy and carbon by
discovery of quinine as a treatment for malaria. In the sixteenth means of photosynthesis, making those resources available not
only for their own needs, but also for the herbivores and omnivores
that consume them, for the carnivores and omnivores that eat the
TABLE 27.1 | Some Medicinal Plants and Their Products herbivores, and for the prokaryotes and fungi that complete food
Plant Product Medical application webs. The earliest steps in human civilization involved cultivating
Belladonna Atropine Dilate pupils for eye examination
angiosperms to provide a reliable food supply.
Pineapple stem Bromelain Control tissue inflammation
Today, twelve species of seed plants account for most of the
Foxglove Digitalin Strengthen heart muscle contraction
food eaten by humans: rice, coconut, wheat, corn (maize), potato,
sweet potato, cassava (also called tapioca or manioc), sugarcane,
Ephedra Ephedrine Ease nasal congestion
sugar beet, soybean, common bean (Phaseolus vulgaris), and ba-
Japanese mint Menthol Relieve coughing
nana. Hundreds of other seed plants are cultivated for food, but
Opium poppy Morphine Relieve pain
none rank with these twelve in importance. Indeed, more than
Pacific yew Paclitaxel Treat ovarian and breast cancers
half of the world’s human population derives the bulk of its food
Cinchona bark Quinine Treat malaria
energy from the seeds of a single plant, Oryza sativa, better known
Curare plant Tubocurarine Muscle relaxant (used in surgery)
as rice. Rice is particularly important in eastern Asia, where it has
Periwinkle Vincristine Treat leukemia and lymphoma
been cultivated for more than 8,000 years (Figure 27.23).

27_Life12e_Ch 27.indd 19 12/4/19 9:42 AM

 KEY CONCEPT
27.4 Recap and Assess
Terrestrial ecosystems could not function without the foods and
habitats provided by plants. Plants produce oxygen and remove
carbon dioxide from the atmosphere, help form soils, and hold
soils in place against erosion by wind and water. Plants also pro-
vide us with many important medicines and are the basis (directly

© Xinhua/Alamy Stock Photo

or indirectly) of virtually all of our food.
1. Name three plants that are used in medicine for humans, and
give an example of how each is used.
2. A parcel of agricultural land that has been used to produce corn for
decades no longer produces healthy crops. Tests indicate that the
soil is depleted of nitrogen and organic matter. A farm extension
Figure 27.23 Rice Feeds Much of the World’s Human Population agent recommends that the farmer grow soybeans and then rotate
this crop with corn in future years. She also recommends that the
These rice fields, or “paddies,” are in south China. Rice has been
farmer till the stubble from the crops after harvest into the soil.
cultivated in this manner for thousands of years.
Why are these practices expected to help crop production?
3. What major group of plants provides (either directly or indirectly) the
In addition to providing food directly for human consumption, majority of calories consumed by humans? Explain your answer.
plants also provide feed for agricultural production of animals. In
particular, the grasses that grow in the world’s rangelands are the As you will see in the coming chapters, the transformation of the
primary forage for the grazing animals that humans use for meat terrestrial environment by plants was critical to the terrestrial in-
and dairy products. vasion by fungi and animals.

▶InvestigatingLIFE

QA
and
How long can most seeds survive,
and why is seed dormancy
important?
Office and handling area
Airlock doors
Sleeve to protect tunnel
from erosion and
climatic changes Tunnel entrance

Courtesy of the Global Crop Diversity Trust

As you saw in Investigating Life: William Beale’s Seed Bridge
Viability Study and in the opening of this chapter, seeds
of some plants can remain dormant but viable for de-
cades, centuries, or in extreme cases, even millennia. Seed vaults
Seed dormancy is especially important for the long-term
survival of plant species that grow in unpredictable
or highly fluctuating environments. If the appropriate
conditions for seed growth and seedling maturation
occur only rarely, then seeds need to be able to remain
Figure 27.24 The Global Seed Vault This vault is located more than 100 meters
dormant in the soil until those favorable conditions oc-
inside a sandstone mountain on the Norwegian island of Spitsbergen, about 1,000 kilo-
cur. Biologists refer to dormant seeds in a given area as
meters from the North Pole. The location was chosen for secure, long-term storage of
the “seed bank.” What conditions will cause the seeds seeds because the surrounding ground was expected to remain perpetually frozen and
to break their dormancy and sprout? Fire, heavy rain, the site is secure against earthquakes, floods, and other natural disasters. However,
appropriate temperatures, or disturbance by animals global warming now threatens the surrounding permafrost.
are some of the more common stimulants for breaking
seed dormancy. Many seeds need to be consumed by an animal and grow the same strain of wheat, for example, and that strain is suscep-
to pass through the acidic conditions of the animal’s digestive sys- tible to a particular pathogen or insect pest, what happens when the
tem before the seeds will sprout. As anyone who has planted a new pathogen or pest spreads and all the susceptible wheat is lost?
garden will know, soil disturbance will often produce the conditions To guard against devastating losses of important plants, many nations
needed for many dormant seeds to start growing. A new garden often around the world have collaborated to develop a secure seed vault,
requires persistent weeding to remove all the unwanted (by the gar- which stores and protects the genetic diversity of crop plants (Figure
dener) plants that have been “awakened” from the seed bank. 27.24). This seed vault is built into a sandstone mountain, which has
long been surrounded by permafrost on the Arctic island of Spitsber-
Future directions gen, Norway (although the permafrost is now threatened by global
We’ve noted that humans depend on a relatively small number of warming). It is situated at a high enough elevation that it will remain
plants for the majority of our food needs, although many more species above sea level even if all the polar ice caps were melted by global
of plants are grown and eaten for food. Traditionally, different locally warming. Cooling systems maintain the vault at –18°C, and even if
adapted strains of crop plants have been grown by different farmers in these systems fail, insulation and local cold weather would protect
different regions of the world. Agricultural companies now produce new the seeds inside and maximize their viability. This seed vault comple-
varieties of crops that increase agricultural production. Unfortunately, ments several shorter-term seed depositories that are maintained in
a downside to these new strains is that farmers across the world have individual countries. Such seed depositories are used by biologists to
switched to growing relatively few varieties of important crop plants, develop new varieties of plant crops and to research the genetic varia-
which increases our global risk to disease and pests. If all farmers tion of plant species from around the world.

27_Life12e_Ch 27.indd 20 12/4/19 9:42 AM

 VISUAL SUMMARY 27
© Zvonimir Atletić/Alamy Stock Photo

You should be able to relate each summary to the adjacent figures. If you go to this visual summary in Achieve,
you can follow links to figures, animations, activities, and simulations that will help you consolidate the material.

KEY CONCEPT Pollen, Seeds, and Wood Contributed

27.1 to the Success of Seed Plants Go to Activity 27.1
■ The two major groups of living seed plants are the gymnosperms and Figure 27.4
angiosperms, both of which include many woody species. A growing pollen tube
■ Seed plants do not require liquid water for fertilization. Pollen grains, Pollen grains
the male gametophytes of seed plants, are carried to a female gametophyte
by wind or by animals.
■ Following pollination, a pollen tube emerges from the pollen grain and
elongates to deliver gametes to the female gametophyte.
■ An ovule consists of the seed plant female gametophyte and the integument
of sporophytic tissue that protects it. The ovule develops into a seed.
Questions © Dr. Jeremy Burgess/Science Source

1. What is the difference Figure 27.5

between pollen and sperm? Unfertilized ovule Fertilized ovule Seed
Seed coat (derived from integument;
2. When you look at a seed plant, parental sporophyte tissue)
which is most obvious—the
haploid gametophyte genera-
tion or the diploid sporophyte Immature female Food supply
(female gametophyte
generation? pine cone
tissue; n)
(cross section) Egg
3. Which parts of a seed plant nucleus (n) Embryo (new
Integument sporophyte; 2n)
constitute the gametophyte
Germinated
generation? Megaspore (n)
pollen grain (n)

Micropyle

Pollen grain (n)

KEY CONCEPT Once Dominant Gymnosperms Still Thrive

27.2 in Some Environments Go to Activity 27.2 and Animation 27.1
■ The gymnosperms produce ovules Figure 27.7
and seeds that are not protected Female cones (megastrobili) in Pinus contorta Male cones (microstrobili) in Pinus contorta
by ovary or fruit tissues.
■
Cross section of Cross section of
a megastrobilus a microstrobilus
The major gymnosperm groups Herbaceous
scale
are the cycads, ginkgos,
Woody Pollen-
gnetophytes, and conifers. scale containing
microspores
Seed
Questions Central
axis
1. Why are forest fires often neces- Central
axis
sary for the long-term survival of
some gymnosperm forests?
2. If gymnosperm seeds are not © Scenics & Science/Alamy
© Scenics & Science/
protected by fruits, then what Alamy Stock Photo
Stock Photo
Male cones,
are the red “berries” on a Female cones, or microstrobili
or megastrobili
juniper tree? © Fritz Pölking/
© Gunter Marx/Alamy Stock Photo

Blickwinkel/AGE Fotostock

27_Life12e_Ch 27.indd 21 12/4/19 9:42 AM

 VISUAL SUMMARY 27
KEY CONCEPT Flowers and Fruits Led to Increased
27.3 Diversification of Angiosperms Go to Animation 27.2

■ Flowers and fruits are unique to the Figure 27.18

angiosperms, distinguishing them from the Amborella
gymnosperms.
■
Common
The floral organs, from the base to the apex ancestor of
of the flower, are the sepals, petals, sta- angiosperms Water lilies
mens, and one or more carpels. Stamens
bear microsporangia in anthers. The carpel
includes an ovary containing ovules and a Star anise
receptive surface called the stigma.
■ Flowers may be pollinated by wind or by
Magnoliids
animals. Many angiosperms have coevolved
with their animal pollinators.
■ The magnoliids are the sister group to the
Monocots
monocots and eudicots (the most species-
rich angiosperm clades).
Questions Eudicots

1. What are three ways that different flowers

attract insect pollinators?
2. Which of the following are true fruits: the
parachute-like structures on a mature dandelion
head; the acorns on an oak tree; a pineapple?

KEY CONCEPT

27.4 Plants Play Critical Roles in Terrestrial Ecosystems

■ Plants provide critical ecosystem services that affect soil, Figure 27.22
water, air quality, and climate.
■ Plants provide humans with many important medicinal
products.
■ Plants are primary producers and as such are the foundation
of terrestrial food webs.

© Calvin Chan/Alamy Stock Photo

Questions
1. What plant produces paclitaxel, an important drug for the
treatment of ovarian and breast cancers?
2. What single plant produces the majority of food energy for
more than half the world’s human population?

Figure 27.23
© Xinhua/Alamy Stock Photo

Life 12e
Oxford University Press
Dragonfly Media Group
Life12e_VS27.18.ai Date 07-10-19
Go to Achieve for the eBook, LearningCurve, animations, activities,
simulations, and additional resources and assignments.

27_Life12e_Ch 27.indd 22 12/4/19 9:42 AM