NATURAL
SELECTION AND EVOLUTION
A. The
Role Of Natural Selection
Evolution is the continuous genetic adaptation of
a population of organisms to its environment over time. Evolution results when
there are changes in genes present in a population. There are three
factors that interact to determine how a species changes over time:
environmental factors that affect organisms, sexual reproduction among the
individuals in the gene pool, and the generation of genetic variety within the
gene pool.
The
idea that some individuals whose gene combinations favor life in their
surroundings will be most likely to survive, reproduce, and pass their genes on
to the next generation is known as the theory of natural selection. The
theory of natural selection was first proposed by Charles Darwin and Alfred
Wallace and was clearly set forth in 1859 by Darwin in his book On the
Origin of Species by Means of Natural Selection, or the Preservation of
Favored Races in the Struggle for Life
B.
What
Influences Natural Selection ?
Spontaneous
mutations are changes in DNA that cannot be tied
to a particular causative agent. It is suspected that cosmic radiation or
naturally occurring mutagenic chemicals might be the cause of many of these
mutations. It is knownthat subjecting organisms to high levels of radiation or
to certain chemicals increases the rate at which mutationsoccur. In order for
mutations to be important in the evolution of organisms, they must be in cells
that will become gametes. Mutations to the cells of the skin or liver will only
affect those specific cells and will not be passed on to the next generation.
A
second very important process involved in generating genetic variety is sexual
reproduction. Although sexual reproduction does not generate new genetic information
the way mutations do, it allows for the recombination of genes into mixtures
that did not occur previously. Each individual entering a population by sexual
reproduction carries a unique combination of genes; approximately half donated
by the mother and half donated by the father. During meiosis, variety is
generated in the gametes through crossing-over between homologous chromosomes
and independent assortment of nonhomologous chromosomes. This results in
millions of possible combinations of genes in the gametes of any individual.
When fertilization occurs, one of the millions of possible sperm unites with
one of the millions of possible eggs, resulting in a genetically unique
individual. The gene mixing that occurs during sexual reproduction is known as genetic
recombination. The new individual has a complete set of genes that is
different from that of any other organism that ever existed.
The
importance of generating new gene combinations is particularly important
because the way genes express themselves in an individual can depend on the
other genes present. Genes don’t always express themselves in the same way. In
order for genes to be selected for or against, they must be expressed in the
phenotype of the individuals possessing them. There are many cases of genes
expressing themselves to different degrees in different individuals. Often the
reason for this difference is unknown. Penetrance is a term used to
describe how often an allele expresses itself when present. Expressivity is
a term used to describe situations in which the gene expresses itself but not
equally in all individuals that have it.
Whenever
a successful organism is examined, it can be shown that it reproduces at a rate
in excess of that necessary to merely replace the parents when they die. Although
the total number of organisms in the species may remain constant, the
individuals that make up the population change. It is this extravagant
reproduction that provides the large surplus of genetically different
individuals that allows natural selection to take place. In fact, to maintain
itself in an ever-changing environment, each species must change in ways that
enhance its ability to adapt to its new environment. For this to occur, members
of the population must be eliminated in a non-random manner. Those individuals
that survive are those that are, for the most part, better suited to the
environment than other individuals. They reproduce more of their kind and
transmit more of their genes to the next generation than do individuals with
genes that do not allow them to be well adapted to the environment in which
they live.
C.
Common
Misunderstandings About Natural Selection
There
are several common misinterpretations associated with the process of natural
selection. The first involves the phrase “survival of the fittest.” Individual
survival is certainly important because those that do not survive will not
reproduce. But the more important factor is the number of descendants an
organism leaves.
Second,
the phrase “struggle for life” does not necessarily refer to open conflict and
fighting. It is usually much more subtle than that. When a resource such as
nesting material, water, sunlight, or food is in short supply, some individuals
survive and reproduce more effectively than others.
A
third common misunderstanding involves significance of phenotypic
characteristics that are not caused by genes. Many organisms survive because
they have characteristics that are not genetically determined. The acquired
characteristics are gained during the life of the organism; they are not
genetically determined and, therefore, cannot be passed on to future
generations through sexual reproduction. Therefore, acquired characteristics
are not important to the processes of natural selection.
D.
Proccess
That Drive Natural Selection
Differential
Survival
As
stated previously, the phrase “survival of the fittest” is often associated
with the theory of natural selection. Although this is recognized as an
oversimplification of the concept, survival is an important factor in
influencing the flow of genes to subsequent generations. If a population
consists of a large number of genetically and phenotypically different
individuals it is likely that some of them will possess characteristics that
make their survival difficult. Therefore, they are likely to die early in life
and not have an opportunity to pass their genes on to the next generation.
Differential
Reproductive Rates
Survival
alone does not always ensure reproductive success. For a variety of reasons,
some organisms may be better able to utilize available resources to produce
offspring. If one individual leaves 100 offspring and another leaves only 2,
the first organism has passed more copies of its genetic information
information on to the next generation than has the second. If we assume that
all 102 individual offspring have similar survival rates, the first organism
has been selected for and its genes have become more common in the subsequent
population.
Differential Mate Selection
Within
animal populations, some individuals may be chosen as mates more n others. This
is called “sexual selection.” Obviously, those that are frequently chosen have
an opportunity to pass on more copies of their genes than those that are rarely
chosen. Characteristics of the more frequently chosen individuals may
involve general characteristics, such as body size or aggressiveness, or
specific conspicuous characteristics attractive to the opposite sex.
E.
Gene-Frequency
Studies and Hardy-Weinberg Equilibrium
In
the early 1900s an English mathematician, G. H. Hardy, and a German physician,
Wilhelm Weinberg, recognized that it was possible to apply a simple
mathematical relationship to the study of gene frequencies. Their basic idea
was that if certain conditions existed, gene frequencies would remain constant,
and that the distribution of genotypes could be described by the relationship A2
+ 2Aa + a2 = 1, where A2 represents the frequency of the
homozygous dominant genotype, 2Aa represents the frequency of the heterozygous
genotype, and a2 represents the frequency of the homozygous recessive
genotype. Constant gene frequencies over several generations would imply that
evolution is not taking place. Changing gene frequencies would indicate
that evolution is taking place.
The
conditions necessary for gene frequencies to remain constant are:
1. Mating must
be completely random.
2. Mutations
must not occur.
3.
Migration of individual organisms into and out of the population must not
occur.
4. The
population must be very large.
5.
All genes must have an equal chance of being passed on to the next generation.
(Natural selection is not occurring.)
The
concept that gene frequencies will remain constant if these five conditions are
met has become known as the Hardy-Weinberg concept. The Hardy-Weinberg
concept is important because it allows a simple comparison of allele frequency
to indicate if genetic changes are occurring. Two different populations of the
same organism can be compared to see if they have the same allele frequencies,
or populations can be examined at different times to see if allele frequencies
are changing.
F.
A
Summary Of The Causes Of Evolutionary Changes
Evolution
was described as the change in gene frequency over time. We can now see that
several different mechanisms operate to bring about this change. Mutations can
either change one allele into another or introduce an entirely new piece of
genetic information into the population. Immigration can introduce new genetic
information if the entering organisms have unique genes. Emigration and death
remove genes from the gene pool. Natural selection systematically filters some
genes from the population, allowing other genes to remain and become more
common. The primary mechanisms involved in natural selection are differences in
deathrates, reproductive rates, and the rate at which individuals are selected
as mates. In addition, gene frequencies are more easily changed in small
populations because events such as death, immigration, emigration, and mutation
can have a greater impact on a small population than on a large population.
2. SPECIATION
AND EVOLUTIONARY CHANGE
A. Species
: A Working Definintion
A species is commonly defined as a population
of organisms whose members have the potential to interbreed naturally to
produce fertile offspring but do not interbreed with other groups. This
is a working definition; it applies in most cases but must be interpreted to
encompass some exceptions. There are two key ideas within this definition.
First, a species is a population of organisms. Second, the definition involves
the ability of individuals within the group to produce fertile offspring.
Obviously, we cannot check every individual to see if it is capable of mating
with any other individual that is similar to it, so we must make some judgment
calls.
B. How
New Species Originate
Geographic Isolation
The geographic area over which a species can be
found is known as its range. The range of the human species is the entire
world. As a species expands its range or environmental conditions change in
some parts of the range, portions of the population can become separated from
the rest. Thus, many species consist of partially isolated populations that
display characteristics significantly different from other local populations.
Many of the differences observed may be directly related to adaptations to
local environmental conditions. This means that new colonies or isolated
populations may have infrequent gene exchange with their geographically distant
relatives.
Speciation is the process of generating new species.
This process has occurred only if gene flow between isolated populations does
not occur even after barriers are removed. the process of speciation can begin
with the geographic isolation of a portion of the species, but new species are
generated only if isolated populations become separate from one another
genetically. Speciation by this method is really a three-step process. It
begins with geographic isolation, is followed by the action of selective agents
that choose specific genetic combinations as being valuable, and ends with the
genetic differences becoming so great that reproduction between the two groups
is impossible.
It is also possible to envision ways in which
speciation could occur without geographic isolation being necessary. Any process
that could result in the reproductive isolation of a portion of a species could
lead to the possibility of speciation.
Polyploidy: Instant Speciation
Polyploidy is a condition of having multiple sets of
chromosomes rather than the normal haploid or diploid number. The increase in
the number of chromosomes can result from abnormal mitosis or meiosis in which
the chromosomes do not separate properly. Because the number of chromosomes of
the polyploid is different from that of the parent, successful reproduction
with the parent species would be difficult. This is because meiosis would
result in gametes that had different chromosome numbers from the original,
parent organism. In one step, the polyploid could be isolated reproductively
from its original species.
C. Maintaining
Genetic Isolation
In order for a new species to continue to exist, it
must reproduce but continue to remain genetically distinct from other similar
species. The speciation process typically involves the development of
reproductive isolating mechanisms or genetic isolating mechanisms. These
mechanisms prevent matings between species and therefore help maintain distinct
species. A great many types of genetic isolating mechanisms are recognized.
D. The
Development Of Evolutionary Thought
In the mid-1700s, Georges-Louis Buffon, a French
naturalist, expressed some curiosity about the possibilities of change
(evolution) in animals, but he did not suggest any mechanism that would result
in evolution. In 1809, Jean-Baptiste de Lamarck, a student of Buffon’s, suggested
a process by which evolution could occur. He proposed that acquired
characteristics could be transmitted to offspring. Although we now know
Lamarck’s theory was wrong (because acquired characteristics are not
inherited), it stimulated further thought as to how evolution could occur.
In 1858, Charles Darwin and Alfred Wallace suggested
the theory of natural selection as a mechanism for evolution. They based their
theory on the following assumptions about
the
nature of living things:
1. All organisms produce more offspring than can
survive.
2. No two organisms are exactly alike.
3. Among organisms, there is a constant struggle for
survival.
4. Individuals that possess favorable
characteristics for their environment have a higher rate of survival and produce
more offspring.
5. Favorable characteristics become more common in
the species, and unfavorable characteristics are lost.
This logic seems simple and obvious today, but
remember that at the time Darwin and Wallace proposed their theory, the
processes of meiosis and fertilization were poorly understood, and the concept
of the gene was only beginning to be discussed. Nearly 50 years after Darwin
and Wallace
suggested
their theory, the rediscovery of the work of Gregor Mendel (chapter 10)
provided an explanation for how characteristics could be transmitted from one
generation to the next.Not only did Mendel’s idea of the gene provide ameans of
passing traits from one generation to the next, it also provided the first step
in understanding mutations, gene flow, and the significance of reproductive
isolation. All of these ideas are interwoven into the modern concept of
evolution.
E. Evolutionary
Patterns Above The Species Level
The basic evolutionary pattern is one of divergent evolution in which individual
speciation events cause successive branches in the evolution of a group of
organisms. Furthermore, it is important to recognize that many extinct species were
very successful organisms for millions of years. They were not failures for
their time but simply did not survive to the present. It is also important to realize
that many currently existing organisms will entually become extinct.
Although divergence is the basic pattern in
evolution, it is possible to superimpose several other patterns on it. One special
evolutionary pattern, characterized by a rapid increase in the number of kinds
of closely related species, is known as adaptive
radiation. Adaptive radiation results in an evolutionary explosion of new species
from a common ancestor. There are basically two situations that are thought to
favor adaptive radiation. One is a
condition in which an organism invades a previously unexploited environment. A second set of conditions that can
favor adaptive radiation is one in which a type of organism evolves a new set
of characteristics that enable it to displace organisms that previously filled
roles in the environment.
Another evolutionary pattern, convergent evolution, occurs when organisms of widely different backgrounds
develop similar characteristics. This particular pattern often leads people to
misinterpret the evolutionary history of organisms.
F. Rates
Of Evolution
When we examine the fossil record, we can often see gradual
changes in physical features of organisms over time. This is such a common
feature of the evolutionary record that biologists refer to this kind of
evolutionary change as gradualism. Charles
Darwin’s view of evolution was based on gradual changes in the features of
specific species he observed in his studies of geology and natural history.
However, as early as the 1940s, some biologists
began to challenge gradualism as the typical model for evolutionary change.
They pointed out that the fossils of some species were virtually unchanged over
millions of years. If gradualism were the only explanation for how species
evolved, then
gradual
changes in the fossil record of a species would always be found. Furthermore,
some organisms appear suddenly in the fossil record and show rapid change from the
time they first appeared.
In 1972, two biologists, Niles Eldredge of the
American Museum of Natural History and Stephen Jay Gould of Harvard University,
proposed the idea of punctuated
equilibrium. This hypothesis suggests that evolution occurs in spurts of
rapid change followed by long periods with little evolutionary change. It is
important to recognize that the punctuated equilibrium concept suggests a
different way of achieving evolutionary change.
G. The
Tentative Nature of The Evolutionary History Of Organisms
It is important to understand that thinking about
the concept of evolution can take us in several different directions. First, it
is clear that genetic changes do occur. Mutations introduce new genes into a
species. This has been demonstrated repeatedly with chemicals and radiation. We
also recognize that species can change. We purposely manipulate the genetic
constitution of our domesticated plants and animals and change their
characteristics to sui our needs. We also recognize that different populations
of the same species show genetic differences. Examination of fossils shows that
species of organisms that once existed are no longer in existence. We can also
demonstrate that new species come into existence. This is easiest to do in
plants with polyploidy. It is clear from this evidence that species are not
fixed, unchanging entities.
However, the fossil record is not a complete record
and new fossils are being discovered every year. There are several reasons why
the fossil record is incomplete. First of all the likelihood that an organism
will become a fossil is low. Most organisms die and decompose leaving no trace
of their existence. In addition, some organisms have very resistant parts that
tend to be preserved while others do not. Clams and insects are abundant in the
fossil record. Worms are not. Finally, the discovery of fossils is often
accidental. It is impossible to search through all the layers of sedimentary
rock on the entire surface of the Earth.
H. Human
Evolution
Various species of Australopithecus and Paranthropus
were present in Africa from about 4.4 million years ago untilabout 1 million
years ago. However, from examining the fossil bones of the leg, pelvis, and
foot, it is apparent that the australopiths were relatively short (males, 1.5
meters or less; females, about 1.1 meters) and stocky and walked upright like
humans.
About 2.5 million years ago the first members of the
genus Homo appeared on the scene. There is considerable disagreement about how
many species there were but Homo habilis is one of the earliest. Homo habilis
had a larger brain (650 cubic centimeters) and smaller teeth than australopiths
and made much more use of stone tools.
About 1.8 million years ago Homo ergaster appeared on
the scene. It was much larger (up to 1.6 meters) than H. habilis (about 1.3
meters) and also had a much larger brain (cranial capacity of 850 cubic
centimeters). A little later a similar species (Homo erectus) appears in the
fossil record.
About 800,000 years ago another hominid, classified as
Homo heidelbergensis, appears in the fossil record. Since fossils of this
species are found in Africa, Europe, and Asia, it appears that they constitute
a second wave of migration of early Homo from Africa to other parts of the
world.
There are two
different theories that seek to explain the origin of Homo sapiens.One theory,
known as the out-of-Africa hypothesis, states that modern humans (Homo sapiens)
originated in Africa as had several other hominid species and migrated from
Africa to Asia and Europe and displaced species such as H. erectus and H. Heidelbergensis
that had migrated into these areas previously. The other theory, known as the
multiregional hypothesis, states that H. erectus evolved into H. sapiens.
During a period of about 1.7 million years, fossils of Homo erectus showed a
progressive increase in the size of the cranial capacity and reduction in the
size of the jaw, so that it becomes difficult to distinguish H. erectus from H.
heidelbergensis and H. Heidelbergensis from H. sapiens. Proponents of this
hypothesis believe that H. heidelbergensis is not a distinct species but an
intermediate between the earlier H. erectus and H. sapiens.
The early evolution of humans has been difficult to
piece together because of the fragmentary evidence. Beginning about 4.4 million
years ago the earliest forms of Australopithecus and Paranthropus showed
upright posture and other humanlike characteristics.
The structure of the jaw and teeth indicates that
the various kinds of australopiths were herbivores. Homo habilis had a larger
brain and appears to have been a scavenger. Several other species of the genus
Homo arose in Africa. These forms appear to have been carnivores. Some of these
migrated to Europe and Asia.The origin of Homo sapiens is in dispute. It may
have arisen inAfrica and migrated throughout the world or evolved from earlier ancestors
found throughout Africa, Asia, and Europe.
3. THE
PLANT BODY
The
plant body creates and maintains an internal environment that differs from the
external environment. Plants accomplish through growth some of the same things
that animals achieve through mobility. they must nevertheless obtain nutrients
not only the raw materials of photosynthesis (carbon dioxide and water), but
also mineral elements such as nitrogen, potassium, and calcium. Seed
plants—even the tallest trees—transport water and minerals from the soil to
their tops, and they transport the products of photosynthesis from the leaves
to their roots and other parts.
a. Vegetative
Organs of the Flowering Plant Body
Flowering
plants (angiosperms) are tracheophytes that are characterized by double
fertilization, a triploid endosperm, and seeds enclosed in modified leaves
called carpels. Their xylem contains cells
called vessel
elements and fibers, and their phloem contains sieve tube elements and
companion cells.
Flowering
plants possess three kinds of vegetative (nonreproductive) organs:
roots, stems, and leaves. Flowers, which are the plant’s devices for sexual
reproduction, consist of modified leaves and stems; flowers will be considered
in detail in a later chapter. Most flowering plants belong to one of two major
lineages. Monocots are generally narrow-leaved flowering plants such as
grasses, lilies, orchids, and palms. Eudicots are broad-leaved flowering
plants such as soybeans, roses, sunflowers, and maples.
In both lineages,
the vegetative plant body consists of two systems: the shoot system and the
root system. The shoot system of a plant consists of the stems, leaves,
and flowers. Broadly speaking, the leaves are the chief
organs of photosynthesis. The stems hold and display
the leaves to the sun and provide connections for the transport of materials
between roots and leaves. The locations where leaves attach to a stem are
called nodes, and
the stem regions between successive nodes are internodes. The
root
system anchors the plant in place and provides nutrition. The extreme
branching of plant roots and their high surface area-to-volume ratio allow them
to absorb water and mineral nutrients from the soil. Each of the vegetative
organs can be understood in terms of its structure. By structure we
mean both its overall form, called its morphology, and
its component cells and tissues and their arrangement, called its anatomy.
There are two principal types of root systems. Many
eudicots have a taproot system: a single,
large, deep-growing primary
root accompanied by
less prominent lateral roots. The
taproot itself often functions
as a nutrient storage organ, as
in carrots. Some plants
have adventitious roots. These roots arise above ground from points
along the stem; some even arise from the leaves.
Unlike roots, stems bear buds of various types. A bud
is an embryonic shoot. A stem bears leaves at its nodes,
and where each leaf meets the stem there is a lateral bud. At
the tip of each stem or branch is an apical bud, which produces
the cells for the upward and outward growth and development
of that shoot.
In gymnosperms and most flowering plants, the leaves
are responsible for most of the plant’s photosynthesis, producing energy-rich
organic molecules and releasing oxygen gas. As photosynthetic organs, leaves
are marvelously adapted for gathering light. Typically, the blade of a
leaf is a thin, flat structure attached to the stem by a stalk called a petiole.
During the daytime, the leaf blade is held by its petiole at an angle almost
perpendicular to the rays of the sun.
b.
Plant
Cells
Plant
cells have all the essential organelles common to eukaryotes they have certain
structures and organelles that distinguish them from many other eukaryotes:
_ They contain chloroplasts
or other plastids.
_ They contain vacuoles.
_ They possess
cellulose-containing cell walls
Cell Wall
The cytokinesis of a plant cell is
completed when the two daughter cells are separated by a cell plate The daughter cells then deposit a gluelike
substance within the cell plate; this substance constitutes the middle lamella.
Next, each daughter cell secretes cellulose and other polysaccharides to form a
primary wall. This deposition and secretion continue as the cell expands to its
final size.Once cell expansion stops, a plant cell may deposit one or more
additional cellulosic layers to form a secondary wall internal to the primary
wall. Secondary walls are often impregnated with unique substances that give
them special properties.
Parenchyma
Cell
The most
numerous cell type in young plants is the parenchyma cell . Parenchyma cells
usually have thin walls, consisting only of a primary wall and the shared middle
lamella.The photosynthetic cells in leaves are parenchyma cells that contain
numerous chloroplasts. In the cytoplasm of these cells, starch is often stored
in specialized plastids called leucoplasts.
Collenchyma
Cell
Collenchyma
cells are supporting cells. Their primary walls are characteristically thick at
the corners of the cells. Collenchyma cells are generally elongated. In these cells,
the primary wall thickens, but no secondary wall forms. Collenchyma provides
support to leaf petioles, nonwoody stems, and growing organs.
Sclerenchyma
Cell
Sclerenchyma
cells have a thickened secondary wall that performs their major function: support.
Many sclerenchyma cells function when dead. There are two types of sclerenchyma
cells: elongated fibers and variously shaped sclereids. Fibers provide
relatively rigid support in wood and other parts of the plant, where they are
often organized into bundles. Sclereids may pack together densely.
Xylem
The xylem of
tracheophytes conducts water from roots to aboveground plant parts. It contains
conducting cells called tracheary elements, which undergo programmed cell death
before they assume their function of transporting water and dissolved minerals.
Phloem
The
transport cells of the phloem, unlike those of the mature xylem, are living
cells. In flowering plants, the characteristic cells of the phloem are sieve
tube elements. These cells meet end-to-end. They form long sieve tubes, which transport
carbohydrates and many other materials from their sources to tissues that
consume or store them. As sieve tube elements mature, plasmodesmata in their end
walls enlarge to form pores, enhancing the connection between neighboring
cells. The result is end walls that look like sieves, called sieve plates. At
functional maturity, a sieve tube element is filled with sieve tube sap,
consisting of water, dissolved sugars, and other solutes
c. Plant Tissues and Tissue Systems.
Vascular plants
have three tissue systems: vascular, dermal, and ground. The vascular tissue system, which includes the xylem and phloem, is
the plant’s plumbing or transport system. All the living cells of the plant
body require a source of energy and chemical building blocks. The dermal tissue system is the outer
covering of the plant. All parts of the young plant body are covered by an
epidermis, which may be a single layer of cells or several layers. The
epidermis contains epidermal cells and may also include specialized cell types,
such as the guard cells that form stomata (pores) in leaves.
The ground tissue
system makes up the rest of the plant. It consists primarily of parenchyma
tissue, often supplemented by collenchyma or sclerenchyma. Ground tissue functions
primarily in storage, support, photosynthesis, and the production of defensive
and attractive substances.
d. Forming the Plant Body
Meristem
There are two types of meristems: Apical
meristems give rise to the primary plant body, which is the entire
body of many plants.Lateral meristems give rise to the secondary
plant body. The stems and roots of some plants (most obviously trees) form
wood and become thick; it is the lateral meristems that give rise to the
tissues responsible for this thickening.
Both root and shoot apical meristems give
rise to a set of cylindrical primary meristems that produce the primary
tissues of the plant body.
These complex tissues are derived from two
lateral meristems: the vascular cambium and the cork cambium. The vascular
cambium is a cylindrical tissue consisting. Without the activity of the cork cambium,
this sloughing off of tissues, including the epidermis, would expose the tree
to potential damage, including excessive water loss or invasion by
microorganisms. The cork cambium produces new protective cells, primarily in
the outwarddirection. The walls of these cork cells become impregnated with
suberin. predominantly of vertically elongated cells that divide frequently.
The
root apical meristem
The root apical meristem produces all the
cells that contribute to growth in the length of the root. Some of the daughter
cells from the apical (tip) end of the root apical meristem contribute to a
root cap, which protects the delicate growing region of the root as it pushes
through the soil. Part of the root apical meristem nearest the tip of the root
forms a quiescent center, in which cell divisions are rare.
The growing region above the apical meristem
comprises the three cylindrical primary meristems: the protoderm, the ground
meristem, and the procambium.
The products of the root’s primary meristems
become root tissues
The protoderm gives rise to the outer layer of cells the epidermis which is adapted for protection of the root
and for the absorption of mineral ions and water. In the zone of maturation,
many of the epidermal cells produce amazingly long, delicate root hairs, which
vastly increase the surface area of the root.
Internal to the epidermis, the ground meristem
gives rise to a region of ground tissue that is many cells thick, called the
cortex. The cells of the cortex are relatively unspecialized and often function
in nutrient storage.
Proceeding inward, we come to the endodermis
of the root, a single cylindrical layer of cells that is the innermost cell
layer of the cortex. Unlike those of other cortical cells, the cell walls of
the endodermal cells contain suberin. The placement of this waterproofing
substance in only certain parts of the cell wall enables the cylindrical ring
of endodermal cells to control the access of water and dissolved ions to the
vascular tissues.
Moving inward past the endodermis, we enter
the vascular cylinder, or stele, produced by the procambium. The stele consists
of three tissues: pericycle, xylem, and phloem
At the very center of the root of a eudicot lies
the xylem—seen in cross section in the shape of a star with a variable number
of points. Between the points are bundles of phloem. In monocots, a region of
parenchyma cells, called the pith, lies in the center of the root. The pith
often stores carbohydrate reserves.
The products of the stem’s primary meristems
become stem tissues
The primary meristems, in turn, give rise to the three tissue systems.
The shoot apical meristem also repetitively lays down the beginnings of leaves
and lateral buds.The growing stem has no protective structure analogous to the
root cap, but the leaf primordia can act as a protective covering for the shoot
apical meristem.
The vascular tissue of a young stem, however, is divided into discrete
vascular bundles. Each vascular bundle contains both xylem and phloem. the stem
contains other important storage and supportive tissues. Internal to the ring
of vascular bundles in eudicots is a storage tissue, the pith, and to the
outside lies a similar storage tissue, the cortex.
Many
stems and roots undergo secondary growth
Secondary growth
results from the activity of the two lateral meristems: vascular cambium and
cork cambium. Initially, the vascular cambium is a single layer of cells lying between
the primary xylem and the primary phloem. The root or stem increases in
diameter when the cells of the vascular cambium divide, producing secondary xylem
cells toward the inside of the root or stem and producing secondary phloem
cells toward the outside.
As the vascular
cambium produces secondary xylem and phloem, its principal cell products are
vessel elements, supportive fibers, and parenchyma cells in the xylem and sieve
tube elements, companion cells, fibers, and parenchyma cells in the phloem.
Cork, cork cambium,
and phelloderm make up the periderm of the secondary plant body. As the vascular
cambium continues to produce secondary vascular tissue, the corky layers are in
turn lost, but the continuous formation of new cork cambia in the underlying
phloem gives rise to new corky layers.
e. Leaf Anatomy Supports Photosynthesis
Most eudicot leaves have two zones of photosynthetic
parenchyma tissue referred to as mesophyll, which means “middle of the
leaf.” Within the mesophyll is a great deal of air space through which carbon
dioxide can diffuse to reach and be absorbed by photosynthesizing cells.
Vascular tissue branches extensively throughout the
leaf, forming a network of veins. Veins extend to within a few cell
diameters of all the cells of the leaf, ensuring that the mesophyll cells are
well supplied with water and minerals. The products of photosynthesis are
loaded into the phloem of the veins for export to the rest of the plant. Guard
cells are modified epidermal cells that change their shape, thereby opening
or closing pores called stomata, which serve as passageways between the environment
and the leaf’s interior. When the stomata are open, carbon dioxide can enter
and oxygen can leave, but water vapor can also be lost.
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