22.1 What Is a Plant?

Traditionally, plants are classified as members of the kingdom Plantae. Plants are eukaryotes that have
cell walls containing cellulose and carry out photosynthesis using chlorophyll a and b. While most plants
are autotrophs, a few are parasites and saprobes (saprophytes, decomposers).

The lives of plants center on the need for sunlight, gas exchange, water, and minerals.

• Plants use the energy from sunlight to photosynthesize. As a result, plants display adaptations
shaped by the need to gather sunlight. Photosynthetic organs such as leaves are typically broad
and flat and are arranged on the stem so as to maximize light absorption.
• Plants require oxygen for cellular respiration and carbon dioxide for photosynthesis. They also
need to release excess oxygen made during photosynthesis. Plants must exchange these gases
with the atmosphere and the soil without losing excessive amounts of water through
evaporation.
• On a sunny day, plants can lose a great deal of water to the air. Water is also used during
photosynthesis when the sun is shining. Thus, land plants have evolved structures that limit
water loss and speed the uptake of water from the ground. As they absorb water, plants also
absorb minerals (naturally occurring chemical compounds). Minerals are nutrients in the soil
needed for plant growth. Many plants have specialized tissues that carry water and nutrients
upward from the soil and distribute the products of photosynthesis throughout the plant body.
Simpler types of plants carry out these functions by diffusion (movement of something due to a
concentration gradient, difference in concentration).

The fossil record indicates that the ancestors of today’s plants were water dwelling organisms similar to
today’s green algae. Most of these photosynthetic eukaryotes were unicellular, although a few were
multicellular. Green algae were first classified as protists, but biologists reclassified them later as plants.
Green algae have cell walls and photosynthetic pigments that are identical to those of plants. They also
have reproductive cycles similar to those of plants. Studies of their genomes suggest that they are so
closely related to other plants that they should be considered part of the plant kingdom.

Fossilized spores of land plants occur in rocks 475 million years old, but there are no fossils of the plants
themselves. The oldest fossils of land plants themselves are found roughly 50 million years later (425
million years old) in the fossil record. Lacking leaves and roots, these plants were only a few centimeters
tall. The greatest challenge that early land plants faced was obtaining water, which they achieved by
growing close to the ground in damp locations. Fossils also suggest that the first true land plants were
still dependent on water to complete their life cycle. Over time, the demands of life on land favored
the evolution of plants more resistant to the drying rays of the sun, more capable of conserving water,
and more capable of reproducing without water. These new land plants changed the environment in
ways that enabled new species to evolve. New ecosystems emerged, and organic matter began to form
soil. Several groups of plants evolved from the first land plants. One group developed into mosses.
Another lineage gave rise to ferns, cone-bearing plants, and flowering plants. These groups have
evolved very different adaptations for a wide range of terrestrial environments.
Botanists divide the plant kingdom into five major groups based on four important features:

• Embryo formation
• Specialized water-conducting tissues
• Seeds
• Flowers

Plants that form embryos are often referred to as “land plants,” even though some of them live in
watery environments. Botanists classify plants into finer groups within these major branches by
comparing the DNA sequences of various species.

The life cycle of land plants has two alternating phases, a diploid (2N) phase and a haploid (N) phase.
The shift between haploid and diploid is known as alternation of generations. The multicellular
diploid (2N) phase is known as sporophyte, or spore-producing plant/phase. The multicellular haploid
(N) phase is known as the gametophyte, or gamete-producing plant/phase.
A sporophyte produces haploid spores through meiosis. These spores grow into multicellular structures
called gametophytes. Each gametophyte produces reproductive cells through mitosis called
gametes—sperm and egg cells. During fertilization, a sperm and egg fuse with each other, producing a
diploid zygote. The zygote develops into a new sporophyte, and the cycle begins again.

An important trend in plant evolution is the reduction in size of the gametophyte and the increasing size
of the sporophyte. Although many green algae do have a diploid sporophyte phase, some do not; their
only multicellular bodies are gametophytes. Mosses and their relatives consist of a relatively large
gametophyte and smaller sporophytes. Ferns and their relatives have a small gametophyte and a larger
sporophyte. Seed plants have an even smaller gametophyte, which is contained within sporophyte
tissues.
 22.2 Seedless Plants
Biologists apply the name “algae” to any photosynthetic eukaryote that is not a land plant. As a result,
algae form a paraphyletic group (not a clade). some algae are classified as protists and others are
classified as plants. Those algae that are grouped with plants are called green algae.

Fossil evidence suggests that the green algae were the first plants, appearing on Earth before plants
emerged on land. The green algae share many characteristics—including their photosynthetic pigments
and cell wall composition—with larger, more complex plants. Green algae are mostly aquatic. They are
found in fresh and salt water, and some moist areas on land. Because most green algae are single cells
or branching filaments, they make direct contact with the water in which they grow. They are able to
absorb moisture and nutrients directly from their surroundings. Therefore, most green algae do not
contain the specialized tissues found in other plants.

Like land plants, many green algae have life cycles that switch back and forth between haploid and
diploid phases. Some green algae may not alternate between haploid and diploid with each and every
generation, however. For example, the single-celled green alga Chlamydomonas can stay in the haploid
stage for multiple generations. As long as living conditions are suitable, the haploid cell reproduces
asexually by mitosis. If environmental conditions become unfavorable, Chlamydomonas can switch to a
stage that reproduces sexually. The cell releases gametes that fuse into a diploid zygote—a sporophyte.
The zygote has a thick protective wall, permitting survival in freezing or drying conditions that would
ordinarily kill it. When conditions once again become favorable, the zygote begins to grow. It divides by
meiosis to produce four flagellated haploid cells. These haploid cells then swim away, mature, and
reproduce asexually.
Many green algae form colonies, providing a hint as to how the first multicellular plants may have
evolved. The freshwater alga, Spirogyra, forms long, threadlike colonies called filaments. Volvox forms
colonies consisting of as few as 500 to as many as 50,000 cells arranged to form hollow spheres. The
cells in a volvox colony are connected to one another by strands of cytoplasm enabling them to
communicate. Although most cells in a Volvox colony are identical, a few gamete-producing cells are
specialized for reproduction. Figuring out how algae made the leap of becoming a multicellular colony
can provide clues to how multicellular organisms such as plants and animals evolved from single cells.

Mosses have a thin waxy layer coating that makes it possible for them to resist drying, and thin
filaments known as rhizoids that anchor them to the ground. Rhizoids also absorb water and minerals
from the surrounding soil.

Mosses belong to a group of plants known as bryophytes. Unlike algae, the bryophytes have specialized
reproductive organs enclosed by other, nonreproductive cells. The bryophytes show a higher degree of
specialization than do the green algae and were among the very first plants to become established on
land. In addition to mosses, the bryophytes include two other groups, known as hornworts and
liverworts. Each of the three groups is generally considered to be a separate phylum.

Most land plants carry water in a specialized tissue called vascular tissue, which contains tubes
hardened with a substance called lignin. Bryophytes, however, do not make lignin and do not contain
true vascular tissue. Bryophytes are small because they lack vascular tissue. They can draw up water no
higher than a meter above the ground. Without strong cell walls hardened by lignin, bryophytes also
cannot support a tall plant body against the pull of gravity. These factors limit the height of bryophytes
and confine them to damp environments.

Like all land plants, bryophytes display alternation of generations. In bryophytes, the gametophyte is the
dominant, recognizable stage of the life cycle. The gametophyte is also the stage that carries out most of
the plant’s photosynthesis. The sporophyte is dependent on the gametophyte for its supply of water and
nutrients. Bryophytes produce sperm cells that swim using flagella. For fertilization to occur successfully,
these sperm cells must be released where there is enough water for them to swim to an egg cell.
Because of this, bryophytes live only in damp habitats where there is standing water for at least part of
the year.
When a moss spore lands in a moist place, it sprouts and grows into a tangled mass of green filaments
(protonema). As this young gametophyte grows, it forms rhizoids that grow into the ground and shoots
(branches) that grow into the air. These shoots grow into the familiar green moss plants. Gametes are
formed in reproductive structures at the tips of the gametophytes. Some bryophyte species produce
both sperm and eggs on the same plant (Monoicy), whereas other species produce sperm and eggs on
separate plants (Dioicy). Eggs are produced in archegonia (singular: archegonium). Sperm are produced
in antheridia (singular: antheridium). Sperm and egg cells fuse to produce a diploid zygote. The zygote
marks the beginning of the sporophyte stage of the life cycle. It develops into a multicellular embryo,
growing within the body of the gametophyte and depending on it for water and nutrients. Eventually,
the sporophyte grows out of the gametophyte and develops a long stalk (seta) ending in a capsule. The
spore capsule is called a sporangium. Inside the capsule, haploid spores are produced by meiosis. When
the capsule ripens, it opens, and haploid spores are scattered to the wind to start the cycle again.

About 420 million years ago, vascular plants emerged. These plants developed true vascular tissue
which enabled them to grow high above the ground. Vascular tissue carries water and nutrients much
more efficiently than does any tissue found in bryophytes.
Vascular plants are known as tracheophytes, after a specialized type of water-conducting cell they
contain. These cells, called tracheids, are hollow tubelike cells with thick cell walls strengthened by
lignin. Tracheids are found in xylem, a tissue that carries water upward from the roots to every part of a
plant. Tracheids are connected end-to-end. Openings between tracheids known as pits allow water to
move through a plant more efficiently than by diffusion alone. Vascular plants also have a second
transport tissue called phloem. Phloem transports solutions of nutrients and carbohydrates produced by
photosynthesis. Like xylem, the main cells of phloem are long and specialized to move fluids throughout
the plant body. Vascular tissue—xylem and phloem—make it possible for vascular plants to move
fluids through their bodies against the force of gravity.

Although all seed-bearing plants are tracheophytes, not all tracheophytes are seed-bearing plants.
Among the seedless vascular plants (pteridophytes) are ferns, horsetails, and club mosses. The most
numerous of which are ferns with more than 11,000 species of ferns living today. Ferns have true
vascular tissue, strong roots, creeping or underground stems called rhizomes, and large leaves called
fronds. Ferns can thrive in areas with little light and are most abundant in wet habitats.

The large plants we recognize as ferns are actually diploid sporophytes. Ferns and other vascular plants
have a life cycle in which the diploid sporophyte is the dominant stage. In the fern life cycle, spores grow
into thin, heart-shaped haploid gametophytes. Although it is tiny, the gametophyte grows
independently (unlike in mosses) of the sporophyte. As in bryophytes, sperm and eggs are produced on
these gametophytes in antheridia and archegonia, respectively. Fertilization requires at least a thin film
of water so that the sperm can swim to the eggs. The diploid zygote produced by fertilization
immediately begins to develop into a new sporophyte plant. As the sporophyte matures, haploid spores
develop on the undersides of the fronds in sporangia, and the cycle begins again.
 22.3 Seed Plants
A characteristic shared by all seed plants is the production of seeds. A seed is a plant embryo and a food
supply, encased in a protective covering. The living plant within a seed is diploid and represents the
early developmental stage of the sporophyte phase of the plant life cycle.

Fossils of seed-bearing plants exist from 360 million years ago. Similarities in DNA sequences from
modern plants provide evidence that today’s seed plants are all descended from common ancestors.
The fossil record indicates that ancestors of seed plants evolved new adaptations that enabled them to
survive in many environments on dry land. Unlike mosses and ferns, the gametes of seed plants do not
need standing water for fertilization. Adaptations that allow seed plants to reproduce without standing
water include a reproductive process that takes place in cones or flowers, the transfer of sperm by
pollination, and the protection of embryos in seeds.

In seed plants, the male gametophytes and the female gametophytes grow and mature directly within
the sporophyte. The gametophytes usually develop in reproductive structures known as cones or
flowers. In fact, seed plants are divided into two groups based on which of these structures they have.
Nearly all gymnosperms bear their seeds directly on the scales of cones. In contrast, flowering plants, or
angiosperms, bear their seeds in flowers inside a layer of tissue that protects the seed. Most flowers
produce both male gametophytes and female gametophytes in each flower. Some species have separate
male and female flowers.

In seed plants, the entire male gametophyte is contained in a tiny structure called a pollen grain. Sperm
produced by this gametophyte do not swim through water to fertilize the eggs. Instead, pollen grains
are carried to the female reproductive structure by wind or animals such as insects. The transfer of
pollen from the male reproductive structure to the female reproductive structure is called pollination.

After fertilization, the zygote contained within a seed grows into a tiny plant—the sporophyte embryo.
The embryo often stops growing while it is still small and contained within the seed. The embryo can
remain in this condition for weeks, months, or even years. A tough seed coat surrounds and protects the
embryo and keeps the contents of the seed from drying out. Seeds can survive long periods of bitter
cold, extreme heat, or drought. The embryo begins to grow when conditions are once again right by
using nutrients from the stored food supply until it can carry out photosynthesis on its own.

Gymnosperms alive today include plants such as cycads, ginkgoes, and conifers which include pines and
firs. Reproduction in conifers takes place in cones, which are produced by the mature sporophyte plant.
Conifers produce two types of cones: pollen cones and seed cones. Pollen cones, also called male cones,
produce the pollen grains. A pollen grain makes up the entire male gametophyte stage of the
gymnosperm life cycle. One of the haploid nuclei in the pollen grain will divide later to produce two
sperm nuclei. The more familiar seed cones, or female cones, produce female gametophytes. Seed
cones are generally much larger than pollen cones. Near the base of each scale of the seed cones are
two ovules, the structures in which the female gametophytes develop. Within the ovules, meiosis
produces haploid cells that grow and divide to produce female gametophytes. These gametophytes may
contain hundreds or thousands of cells. When mature, each gametophyte contains a few large egg cells,
each ready for fertilization by sperm nuclei.
The conifer life cycle typically takes two years to complete. The life cycle of pines, for example, begins in
the spring as male cones release enormous numbers of pollen grains that are carried away by the wind.
Some of these pollen grains reach female cones. There, pollen grains are caught in a sticky secretion on
the scales of the female cone and pulled inside toward the ovule. In gymnosperms, the direct transfer
of pollen to the female cone allows fertilization to take place without the need for gametes to swim
through standing water.

If a pollen grain lands near an ovule, the grain splits open and begins to grow a structure called a pollen
tube, which contains two haploid sperm nuclei. Once the pollen tube reaches the newly developed
female gametophyte, one sperm nucleus disintegrates; the other fertilizes the egg contained within the
female gametophyte. Fertilization produces a diploid zygote, which grows into an embryo—the new
sporophyte plant. The embryo is then encased to form a seed. The seed is ready to be scattered by the
wind and grow into a new plant.
 22.4 Flowering Plants
Flowering plants, or angiosperms, first appeared during the Cretaceous Period, about 135 million years
ago, making their origin the most recent of all plant phyla. Flowering plants originated on land and soon
came to dominate Earth’s plant life. Angiosperms make up the vast majority of plant species.
Angiosperms develop unique reproductive organs known as flowers. Flowers contain ovaries, which
surround and protect the seeds. Angiosperms reproduce sexually by means of flowers. After
fertilization, ovaries within flowers develop into fruits that surround, protect, and help disperse the
seeds.

In general, flowers are an evolutionary advantage to plants because they attract animals such as insects
and birds. These animals—drawn by the color, scent, or even the shape of the flower—carry pollen with
them as they leave. Because these animals go directly from flower to flower, they can carry pollen to the
next flower they visit. This means of pollination is much more efficient than the wind pollination of most
gymnosperms.

After pollination, the ovary develops into a fruit. The angiosperm fruit is a structure containing one or
more matured ovaries. The wall of the fruit helps disperse the seeds inside it, carrying them away from
the parent plant. Consider what happens when an animal eats a fleshy fruit, such as a berry. Seeds from
the fruit generally enter the animal’s digestive system. By the time these seeds leave the digestive
system—ready to sprout—the animal may have traveled many kilometers. By using fruit, flowering
plants increase the ranges they inhabit. The fruit—a unique feature of angiosperms—is yet another
reason for their success.

The great diversity of angiosperms has made them especially difficult to classify scientifically. For many
years, flowering plants were classified according to the number of seed leaves, or cotyledons, in their
embryos. Those with one seed leaf were called monocots. Those with two seed leaves were called
dicots. At one time, these two groups were considered classes within the angiosperm phylum, and all
angiosperms were placed in one class or the other. Recent studies of plant genomes and new fossil
discoveries have shown that things are more complicated than that. For example, in 2002, a plant fossil
was discovered in China. Given the name Archaefructus, this organism is the oldest known plant with
reproductive organs like those found in modern flowers. It is more ancient than modern-day monocots
and dicots and can’t be classified as either. Other recent evidence suggests that Amborella, another
plant, belongs to still another ancient lineage of plants. Information gained from the Amborella
discovery led scientists to place other plants, such as the water lilies, near the base of angiosperm
evolution. Scientific classification now places the monocots into a single group but places the dicots in a
variety of distinct and different categories. This means, of course, that the term dicot is no longer used
for classification. However, it can still be used to describe many of the characteristics of plant structure,
and that is how it is used in this book.
Angiosperms are often grouped according to the number of their seed leaves, the strength and
composition of their stems, and the number of growing seasons they live. Naturally, these categories
overlap.

Although we no longer classify monocots and dicots as scientific groups, the terms are still useful.
Monocots and dicots differ in characteristics such as the distribution of vascular tissue in stems, roots,
and leaves, and the number of petals per flower. Monocots include plants such as corn, wheat, lilies,
orchids, and palms. Monocot grasses—especially wheat, corn, and rice—have the important distinction
of being the first plants to be cultivated in mass quantities for food. Dicots include roses, clover,
tomatoes, oaks, and daisies.

The flowering plants can also be subdivided into groups according to the characteristics of their stems.
One of the most important and noticeable stem characteristics is woodiness. Woody plants are made
primarily of cells with thick cell walls that support the plant body. Woody plants include trees, shrubs,
and vines. Shrubs are typically smaller than trees, and vines have stems that are long and flexible. Plant
stems that are smooth and nonwoody are characteristic of herbaceous plants. Herbaceous plants do
not produce wood as they grow. Examples of herbaceous plants include dandelions, zinnias, petunias,
and sunflowers.
Plants can be categorized according to their life spans. Some plants grow, flower, and die in a single
year. Other types of plants continue to grow from year to year. The life span of plants is determined by a
combination of genetic and environmental factors.