Plant - biology.

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Plant - biology.
I

INTRODUCTION

Plant, any member of the plant kingdom, comprising about 260,000 known species of mosses, liverworts, ferns, herbaceous and woody plants, bushes, vines, trees,
and various other forms that mantle the Earth and are also found in its waters. Plants range in size and complexity from small, nonvascular mosses, which depend on
direct contact with surface water, to giant sequoia trees, which can draw water and minerals through their vascular systems to elevations of more than 100 m (330 ft).
Only a tiny percentage of plant species are directly used by humans for food, shelter, fiber, and drugs. At the head of the list are rice, wheat, corn, legumes, cotton,
conifers, and tobacco, on which whole economies and nations depend. Of even greater importance to humans are the indirect benefits reaped from the entire plant
kingdom and its more than 1 billion years of carrying out photosynthesis. Plants have laid down the fossil fuels that provide power for industrial society, and throughout
their long history plants have supplied sufficient oxygen to the atmosphere to support the evolution of higher animals. Today the world's biomass is composed
overwhelmingly of plants, which not only underpin almost all food webs, but also modify climates and create and hold down soil, making what would otherwise be stony,
sandy masses habitable for life.

II

DIFFERENTIATION FROM OTHER KINGDOMS

Plants are multicellular eukaryotes--that is, their cells contain membrane-bound structures called organelles. Plants differ from other eukaryotes because their cells are
enclosed by more or less rigid cell walls composed primarily of cellulose. The most important characteristic of plants is their ability to photosynthesize. During
photosynthesis, plants make their own food by converting light energy into chemical energy--a process carried out in the green cellular organelles called chloroplasts
(see Chlorophyll; Chloroplast). A few plants have lost their chlorophyll and have become saprophytes or parasites--that is, they absorb their food from dead organic
matter or living organic matter, respectively--but details of their structure show that they are evolved plant forms.
Fungi, also eukaryotic and long considered members of the plant kingdom, have now been placed in a separate kingdom because they lack chlorophyll and plastids and
because their rigid cell walls contain chitin rather than cellulose. Unlike the majority of plants, fungi do not manufacture their own food; instead they are saprophytic,
absorbing their food from either dead or living organic matter.
The various groups of algae were also formerly placed in the plant kingdom because many are eukaryotic and because most have rigid cell walls and carry out
photosynthesis. Nonetheless, because of the variety of pigment types, cell wall types, and physical attributes found in the algae, they are now recognized as part of two
separate kingdoms, containing a diversity of plantlike and other organisms that are not necessarily closely related. One of the phyla of algae, the green algae, is
believed to have given rise to the plant kingdom, because its chlorophylls, cell walls, and other details of cellular structure are similar to those of plants.
The animal kingdom is also multicellular and eukaryotic, but its members differ from the plants in deriving nutrition from other organic matter; by ingesting food rather
than absorbing it, as in the fungi; by lacking rigid cell walls; and, usually, by having sensory capabilities and being motile, at least at some stage. See Classification.

III

PLANT PHYLA

The many species of organisms in the plant kingdom are divided into several phyla, or divisions, totaling about 260,000 species. The bryophytes are a diverse
assemblage of three phyla of nonvascular plants, with about 16,000 species, that includes the mosses, liverworts, and hornworts. Bryophytes lack a well-developed
vascular system for the internal conduction of water and nutrients and have been called nonvascular plants. It takes two generations to complete the plant life cycle
(Alternation of Generations). The familiar leafy plant of bryophytes is the sexual, or gamete-producing, generation of the life cycle of these organisms. Because of the
lack of a vascular system and because the gametes require a film of water for dispersal, bryophytes are generally small plants that tend to occur in moist conditions,
although some attain large size under favorable circumstances and others (usually very small) are adapted to desert life.
The other phyla are collectively termed vascular plants, or tracheophytes. Vascular tissue is internal conducting tissue for the movement of water, minerals, and food.
There are two types of vascular tissue: xylem, which conducts water and minerals from the ground to stems and leaves, and phloem, which conducts food produced in
the leaves to the stems, roots, and storage and reproductive organs. Besides the presence of vascular tissue, tracheophytes contrast with bryophytes in that
tracheophyte leafy plants are the asexual, or spore-producing, generation of their life cycle. In the evolution of tracheophytes, the spore-producing generation became
much larger and more complex, whereas the gamete-producing generation became reduced and merely contained in the sporophyte tissue. This ability to evolve into
larger and more diverse sporophytes, together with the ability of the vascular system to elevate water, freed tracheophytes from direct dependence on surface water.
They were thus able to dominate all the terrestrial habitats of the Earth, except the higher Arctic zones, and to provide food and shelter for its diverse animal
inhabitants.

IV

CELL STRUCTURE AND FUNCTION

The tremendous variety of plant species is, in part, a reflection of the many distinct cell types that make up individual plants. Fundamental similarities exist among all
these cell types, however, and these similarities indicate the common origin and the interrelationships of the different plant species. Each individual plant cell is at least
partly self-sufficient, being isolated from its neighbors by a cell membrane, or plasma membrane, and a cell wall. The membrane and wall allow the individual cell to
carry out its functions; at the same time, communication with surrounding cells is made possible through cytoplasmic connections called plasmodesmata.

A

Cell Wall

The most important feature distinguishing the cells of plants from those of animals is the cell wall. In plants this wall protects the cellular contents and limits cell size. It
also has important structural and physiological roles in the life of the plant, being involved in transport, absorption, and secretion.
A plant's cell wall is composed of several chemicals, of which cellulose (made up of molecules of the sugar glucose) is the most important. Cellulose molecules are united
into fibrils, which form the structural framework of the wall. Other important constituents of many cell walls are lignins, which add rigidity, and waxes, such as cutin and
suberin, which reduce water loss from cells. Many plant cells produce both a primary cell wall, while the cell is growing, and a secondary cell wall, laid down inside the
primary wall after growth has ceased. Plasmodesmata penetrate both primary and secondary cell walls, providing pathways for transporting substances.

B

Protoplast

Within the cell wall are the living contents of the cell, called the protoplast. These contents are bounded by a cell membrane composed of a phospholipid bi-layer. The
protoplast contains the cytoplasm, which in turn contains various membrane-bound organelles and vacuoles and the nucleus, which is the hereditary unit of the cell.

B1

Vacuoles

Vacuoles are membrane-bound cavities filled with cell sap, which is made up mostly of water containing various dissolved sugars, salts, and other chemicals.

B2

Plastids

Plastids are types of organelles, structures that carry out specialized functions in the cell. Three kinds of plastids are important here. Chloroplasts contain chlorophylls
and carotenoid pigments; they are the site of photosynthesis, the process in which light energy from the sun is fixed as chemical energy in the bonds of various carbon
compounds. Leucoplasts, which contain no pigments, are involved in the synthesis of starch, oils, and proteins. Chromoplasts manufacture carotenoids.

B3

Mitochondria

Whereas plastids are involved in various ways in storing energy, another class of organelles, the mitochondria, are the sites of cellular respiration. This process involves
the transfer of chemical energy from carbon-containing compounds to adenosine triphosphate, or ATP, the chief energy source for cells. The transfer takes place in
three stages: glycolysis (in which acids are produced from carbohydrates); the Krebs cycle, also called the citric acid cycle; and electron transfer. Like plastids,
mitochondria are bounded by two membranes, of which the inner one is extensively folded; the folds serve as the surfaces on which the respiratory reactions take
place.

B4

Ribosomes, Golgi Apparatus, and Endoplasmic Reticulum

Two other important cellular contents are the ribosomes, the sites at which amino acids are linked together to form proteins, and the Golgi apparatus, which plays a role
in the secretion of materials from cells. In addition, a complex membrane system called the endoplasmic reticulum runs through much of the cytoplasm and appears to
function as a communication system; various kinds of cellular substances are channeled through it from place to place. Ribosomes are often connected to the
endoplasmic reticulum, which is continuous with the double membrane surrounding the nucleus of the cell.

B5

Nucleus

The nucleus controls the ongoing functions of the cell by specifying which proteins are produced. It also stores and passes on genetic information to future generations
of cells during cell division. See Cell.

V

TISSUE SYSTEMS

There are many variants of the generalized plant cell and its parts. Similar kinds of cells are organized into structural and functional units, or tissues, which make up the
plant as a whole, and new cells (and tissues) are formed at growing points of actively dividing cells. These growing points, called meristems, are located either at the
stem and root tips (apical meristems), where they are responsible for the primary growth of plants, or laterally in stems and roots (lateral meristems), where they are
responsible for secondary plant growth. Three tissue systems are recognized in vascular plants: dermal, vascular, and ground (or fundamental).

A

Dermal System

The dermal system consists of the epidermis, or outermost layer, of the plant body. It forms the skin of the plant, covering the leaves, flowers, roots, fruits, and seeds.
Epidermal cells vary greatly in function and structure.
The epidermis may contain stomata, openings through which gases are exchanged with the atmosphere. These openings are surrounded by specialized cells called
guard cells, which, through changes in their size and shape, alter the size of the stomatal openings and thus regulate the gas exchange. The epidermis is covered with a
waxy coating called the cuticle, which functions as a waterproofing layer and thus reduces water loss from the plant surface through evaporation. If the plant undergoes
secondary growth--growth that increases the diameter of roots and stems through the activity of lateral meristems--the epidermis is replaced by a peridermis made up
of heavily waterproofed cells (mainly cork tissue) that are dead at maturity.

B

Vascular System

The vascular tissue system consists of two kinds of conducting tissues: the xylem, responsible for conduction of water and dissolved mineral nutrients, and the phloem,
responsible for conduction of food. The xylem also stores food and helps support the plant.

B1

Xylem

The xylem consists of two types of conducting cells: tracheids and vessels. Elongated cells, with tapered ends and secondary walls, both types lack cytoplasm and are
dead at maturity. The walls have pits--areas in which secondary thickening does not occur--through which water moves from cell to cell. Vessels usually are shorter and
broader than tracheids, and in addition to pits they have perforation--areas of the cell wall that lack both primary and secondary thickenings and through which water
and dissolved nutrients may freely pass.

B2

Phloem

The phloem, or food-conducting tissue, consists of cells that are living at maturity. The principal cells of phloem, the sieve elements, are so called because of the clusters
of pores in their walls through which the protoplasts of adjoining cells are connected. Two types of sieve elements occur: sieve cells, with narrow pores in rather uniform
clusters on the cell walls, and sieve-tube members, with larger pores on some walls of the cell than on others. Although the sieve elements contain cytoplasm at
maturity, the nucleus and other organelles are lacking. Associated with the sieve elements are companion cells that do contain nuclei and that are responsible for
manufacturing and secreting substances into the sieve elements and removing waste products from them.

C

Ground System

The ground, or fundamental, tissue systems of plants consist of three types of tissue. The first, called parenchyma, is found throughout the plant and is living and
capable of cell division at maturity. Usually only primary walls are present, and these are uniformly thickened. The cells of parenchyma tissue carry out many specialized

physiological functions--for example, photosynthesis, storage, secretion, and wound healing. They also occur in the xylem and phloem tissues.
Collenchyma, the second type of ground tissue, is also living at maturity and is made up of cells with unevenly thickened primary cell walls. Collenchyma tissue is pliable
and functions as support tissue in young, growing portions of plants.
Sclerenchyma tissue, the third type, consists of cells that lack protoplasts at maturity and that have thick secondary walls usually containing lignin. Sclerenchyma tissue
is important in supporting and strengthening those portions of plants that have finished growing.

VI

PLANT ORGANS

The body of a vascular plant is organized into three general kinds of organs: roots, stems, and leaves. These organs all contain the three kinds of tissue systems
mentioned above, but they differ in the way the cells are specialized to carry out different functions.

A

Roots

The function of roots is to anchor the plant to its substrate and to absorb water and minerals. Thus, roots are generally found underground and grow downward, or in
the direction of gravity. Unlike stems, they have no leaves or nodes. The epidermis is just behind the growing tip of roots and is covered with root hairs, which are
outgrowths of the epidermal cells. The root hairs increase the surface area of the roots and serve as the surface through which water and nutrients are absorbed.
Internally, roots consist largely of xylem and phloem, although many are highly modified to carry out specialized functions. Thus, some roots are important food and
storage organs--for example, beets, carrots, and radishes. Such roots have an abundance of parenchyma tissue. Many tropical trees have aerial prop roots that serve
to hold the stem in an upright position. Epiphytes have roots modified for quick absorption of rainwater that flows over the bark of the host plants.
Roots increase in length through the activity of apical meristems and in diameter through the activity of lateral meristems. Branch roots originate internally at some
distance behind the growing tip, when certain cells become meristematic.

B

Stems

Stems usually are above ground, grow upward, and bear leaves, which are attached in a regular pattern at nodes along the stem. The portions of the stem between
nodes are called internodes. Stems increase in length through the activity of an apical meristem at the stem tip. This growing point also gives rise to new leaves, which
surround and protect the stem tip, or apical bud, before they expand. Apical buds of deciduous trees, which lose their leaves during part of the year, are usually
protected by modified leaves called bud scales.
Stems are more variable in external appearance and internal structure than are roots, but they also consist of the three tissue systems and have several features in
common. Vascular tissue is present in bundles that run the length of the stem, forming a continuous network with the vascular tissue in the leaves and the roots. The
vascular tissue of herbaceous plants is surrounded by parenchyma tissue, whereas the stems of woody plants consist mostly of hard xylem tissue. Stems increase in
diameter through the activity of lateral meristems, which produce the bark and wood in woody plants. The bark, which also contains the phloem, serves as a protective
outer covering, preventing damage and water loss.
Within the plant kingdom are many modifications of the basic stem, such as the thorns of hawthorns. Climbing stems, such as the tendrils of grapes and Boston ivy,
have special modifications that allow them to grow up and attach to their substrate. Many plants, such as cacti, have reduced leaves or no leaves at all, and their stems
act as the photosynthetic surface. Some stems, including those of many grasses, creep along the surface of the ground and create new plants through a process called
vegetative reproduction. Other stems are borne underground and serve as food-storage organs, often allowing the plant to survive through the winter; the so-called
bulbs of the tulip and the crocus are examples.

C

Leaves

The leaf is the primary photosynthetic organ of most plants. Leaves are usually flattened blades that consist, internally, mostly of parenchyma tissue called the
mesophyll, which is made up of loosely arranged cells with spaces between them. The spaces are filled with air, from which the cells absorb carbon dioxide and into
which they expel oxygen. The mesophyll is bounded by the upper and lower surface of the leaf blade, which is covered by epidermal tissue. A vascular network runs
through the mesophyll, providing the cell walls with water and removing the food products of photosynthesis to other parts of the plants.
The leaf blade is connected to the stem through a narrowed portion called the petiole, or stalk, which consists mostly of vascular tissue. Appendages called stipules are
often present at the base of the petiole.
Many specialized forms of leaves occur. Some are modified as spines, which help protect plants from predators. Insectivorous plants possess highly modified leaves that
trap and digest insects to obtain needed nutrients. Some leaves are brightly colored and petal-like, serving to attract pollinators to otherwise small, unattractive flowers.
Perhaps the most highly modified leaves are flowers themselves. The individual parts of flowers--carpels, stamens, petals, and sepals--are all modified leaves that have
taken on reproductive functions.

VII

GROWTH AND DIFFERENTIATION

The growth and differentiation of the various plant tissue and organ systems are controlled by various internal and external factors.

A

Hormones

Plant hormones, specialized chemical substances produced by plants, are the main internal factors controlling growth and development. Hormones are produced in one
part of a plant and transported to others, where they are effective in very small amounts. Depending on the target tissue, a given hormone may have different effects.
Thus, auxin, one of the most important plant hormones, is produced by growing stem tips and transported to other areas where it may either promote growth or inhibit
it. In stems, for example, auxin promotes cell elongation and the differentiation of vascular tissue, whereas in roots it inhibits growth in the main system but promotes
the formation of adventitious roots. It also retards the abscission (dropping off) of flowers, fruits, and leaves.
Gibberellins are other important plant-growth hormones; more than 50 kinds are known. They control the elongation of stems, and they cause the germination of some
grass seeds by initiating the production of enzymes that break down starch into sugars to nourish the plant embryo. Cytokinins promote the growth of lateral buds,
acting in opposition to auxin; they also promote bud formation. In addition, plants produce the gas ethylene through the partial decomposition of certain hydrocarbons,
and ethylene in turn regulates fruit maturation and abscission.

B

Tropisms

Various external factors, often acting together with hormones, are also important in plant growth and development. One important class of responses to external stimuli
is that of the tropisms--responses that cause a change in the direction of a plant's growth. Examples are phototropism, the bending of a stem toward light, and
geotropism, the response of a stem or root to gravity. Stems are negatively geotropic, growing away from gravity, whereas roots are positively geotropic.
Photoperiodism, the response to 24-hour cycles of dark and light, is particularly important in the initiation of flowering. Some plants are short-day, flowering only when
periods of light are less than a certain length (see Biological Clocks). Other variables--both internal, such as the age of the plant, and external, such as
temperature--are also involved with the complex beginnings of flowering.

VIII

ECOLOGY

Rooted as they are in the ground, plants are commonly thought of as leading sedentary, vegetative, passive lives. However, a look at the ingeniously developed
interactions that plants have with the other organisms in their ecosystems quickly corrects this notion.

A

Cooperation and Competition

Many plant species exist as separate male and female plants, and pollen from male flowers must reach the female flowers in order for pollination and seed development
to take place. The agent of pollination is sometimes the wind (a part of the physical environment), but in many cases it is an insect, bat, or bird (members of the
biological environment). Plants may also rely on agents for dispersing their seed. Thus, after pollination, cherry trees develop cherries that attract birds, which ingest
the fruit and excrete the cherry stones in more distant terrains.
Plants have evolved many other mutually beneficial relationships, such as the nitrogen-fixing bacteria that occur in the nodules on the roots of legumes (see Nitrogen
Fixation). Many prairie grasses and other plants that flourish on open land depend on various herbivores to keep forests from closing in and shading them.
In the competition among plants for light, many species have evolved such mechanisms as leaf shape, crown shape, and increased height in order to intercept the sun's
rays. In addition, many plants produce chemical substances that inhibit the germination or establishment of seeds of other species near them, thus excluding competing
species from mineral resources as well as light. Walnut species, for example, use such an allelopathy, or chemical inhibition.

B

The Food Web

Because plants are autotrophs--organisms that are able to manufacture their own food--they lie at the very foundation of the food web. Heterotrophs--organisms that
cannot manufacture their own food--usually lead less sedentary lives than plants, but they ultimately depend on autotrophs as sources of food. Plants are first fed upon
by primary consumers, or herbivores, which in turn are fed upon by secondary consumers, or carnivores. Decomposers act upon all levels of the food web. A large
portion of energy is lost at each step in the food web; only about 10 percent of the energy in one level is stored by the next. Thus, most food webs contain only a few
steps.

C

Plants and Humans

From the prehistoric beginnings of agriculture until recent times, only a few of the total plant species have been taken from the wild and refined to become primary
sources of food, fiber, shelter, and drugs. This process of plant cultivation and breeding began largely by accident, possibly as the seeds of wild fruits and vegetables,
gathered near human habitations, sprouted and were crudely cultivated. Plants such as wheat, which possibly originated in the eastern Mediterranean region more than
9,000 years ago, were selected and replanted year after year for their superior food value; today many domesticated plants can scarcely be traced back to their wild
ancestors or to the original plant communities in which they originated. This selective process took place with no prior knowledge of plant breeding but, rather, through
the constant and close familiarity that preindustrial humans had with plants.
Today, however, the human relationship with plants is nearly reversed: An increasing majority of people have little or no contact with plant cultivation, and the farmers
that do have such contact are becoming more and more specialized in single crops. The breeding process, on the other hand, has been greatly accelerated, largely
through advances in genetics. Plant geneticists are now able to develop, in only a few years, such plant strains as wind-resistant corn, thus greatly increasing crop
yields.
At the same time, humans have accelerated the demand for food and energy to the extent that entire species and ecosystems of plants are being destroyed before
scientists can develop an understanding of which plant species have the potential to benefit humanity. Most species remain little known; those that seem to offer the
greatest hope for providing new sources of food, drugs, and other useful products exist in tropical rain forests and other areas where rapidly growing human
populations can quickly reduce the land to arid, sandy wastes. According to the World Conservation Union, about 34,000 species of plants are at risk of becoming
extinct. This amounts to about one of every eight known species of ferns, flowering plants, and conifers and related plants. Increased knowledge of plants and attention
to their survival are needed to solve many of the problems confronting the human world today. (World Food Supply.)
See also Dicots; Diseases of Plants; Fruit; Monocots; Nut; Plant Distribution; Plant Propagation; and Poisonous Plants.

Contributed By:
Marshall R. Crosby
Peter H. Raven
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