Module 4
Module 4
During the vegetative phase of growth, plants increase in size and produce a shoot system
and a root system. As they enter the reproductive phase, some of the branches start to bear
flowers. Many flowers are borne singly, whereas some are borne in clusters. The flower is borne
on a stalk known as a receptacle. Flower shape, color, and size are unique to each species, and
are often used by taxonomists to classify plants.
If all four whorls (the calyx, corolla, androecium, and gynoecium) are present, the flower
is described as complete. If any of the four parts is missing, the flower is known as incomplete.
Flowers that contain both an androecium and a gynoecium are called perfect, androgynous or
hermaphrodites. There are two types of incomplete flowers: staminate flowers contain only an
androecium, and carpellate flowers have only a gynoecium (figure 3).
Figure 3. The corn plant has both staminate (male)
and carpellate (female) flowers. Staminate flowers,
which are clustered in the tassel at the tip of the
stem, produce pollen grains. Carpellate flower are
clustered in the immature ears. Each strand or silk
is a stigma. The corn kernel are seeds that develop
on the ear after fertilization. Also shown is the
lower stem and root.
If both male and female flowers are borne on the same plant, the species is called
monoecious (meaning “one home”): examples are corn and pea. Species with male and female
flowers borne on separate plants are termed dioecious, or “two homes,” examples of which
are C. papaya and Cannabis. The ovary, which may contain one or multiple ovules, may be
placed above other flower parts, which is referred to as superior; or, it may be placed below the
other flower parts, referred to as inferior (figure 4).
Figure 4. The (a) lily is a superior flower, which has the ovary above the
other flower parts. (b) Fuchsia is an inferior flower, which has the ovary
beneath other flower parts.
Figure 5. Shown is (a) a cross section of an anther at two developmental stages. The
immature anther (top) contains four microsporangia, or pollen sacs. Each microsporangium
contains hundreds of microspore mother cells that will each give rise to four pollen grains.
The tapetum supports the development and maturation of the pollen grains. Upon maturation
of the pollen (bottom), the pollen sac walls split open and the pollen grains (male
gametophytes) are released. (b) In these scanning electron micrographs, pollen sacs are
ready to burst, releasing their grains.
Within the microsporangium, the microspore mother cell divides by meiosis to give rise
to four microspores, each of which will ultimately form a pollen grain (figure 6). An inner layer
of cells, known as the tapetum, provides nutrition to the developing microspores and contributes
key components to the pollen wall. Mature pollen grains contain two cells: a generative cell and
a pollen tube cell. The generative cell is contained within the larger pollen tube cell. Upon
germination, the tube cell forms the pollen tube through which the generative cell migrates to
enter the ovary. During its transit inside the pollen tube, the generative cell divides to form two
male gametes (sperm cells). Upon maturity, the microsporangia burst, releasing the pollen grains
from the anther.
Each pollen grain has two coverings: the exine (thicker, outer layer) and the intine. The
exine contains sporopollenin, a complex waterproofing substance supplied by the tapetal cells.
Sporopollenin allows the pollen to survive under unfavorable conditions and to be carried by
wind, water, or biological agents without undergoing damage.
Female Gametophyte (The Embryo Sac)
A double-layered integument protects the megasporangium and, later, the embryo sac.
The integument will develop into the seed coat after fertilization and protect the entire seed. The
ovule wall will become part of the fruit. The integuments, while protecting the megasporangium,
do not enclose it completely, but leave an opening called the micropyle. The micropyle allows
the pollen tube to enter the female gametophyte for fertilization.
Figure 8. This image shows the life cycle of a conifer. Pollen from male
cones blows up into upper branches, where it fertilizes female cones.
Examples are shown of female and male cones.
Male Gametophyte
A male cone has a central axis on which bracts, a type of modified leaf, are attached. The bracts
are known as microsporophyll (figure 9) and are the sites where microspores will develop. The
microspores develop inside the microsporangium. Within the microsporangium, cells known as
microsporocytes divide by meiosis to produce four haploid microspores. Further mitosis of the
microspore produces two nuclei: the generative nucleus, and the tube nucleus. Upon maturity,
the male gametophyte (pollen) is released from the male cones and is carried by the wind to land
on the female cone.
Female Gametophyte
The female cone also has a central axis on which bracts known as megasporophylls (figure 9)
are present. In the female cone, megaspore mother cells are present in the megasporangium. The
megaspore mother cell divides by meiosis to produce four haploid megaspores. One of the
megaspores divides to form the multicellular female gametophyte, while the others divide to
form the rest of the structure. The female gametophyte is contained within a structure called the
archegonium.
Figure 9. These series of micrographs show male and female gymnosperm gametophytes. (a) This male
cone, shown in cross section, has approximately 20 microsporophyll, each of which produces hundreds of
male gametophytes (pollen grains). (b) Pollen grains are visible in this single microsporophyll. (c) This
micrograph shows an individual pollen grain. (d) This cross section of a female cone shows portions of
about 15 megasporophylls. (e) The ovule can be seen in this single megasporophyll. (f) Within this single
ovule are the megaspore mother cell (MMC), micropyle, and a pollen grain.
Reproductive Process
Upon landing on the female cone, the tube cell of the pollen forms the pollen tube,
through which the generative cell migrates towards the female gametophyte through the
micropyle. It takes approximately one year for the pollen tube to grow and migrate towards the
female gametophyte. The male gametophyte containing the generative cell splits into two sperm
nuclei, one of which fuses with the egg, while the other degenerates. After fertilization of the
egg, the diploid zygote is formed, which divides by mitosis to form the embryo. The scales of the
cones are closed during development of the seed. The seed is covered by a seed coat, which is
derived from the female sporophyte. Seed development takes another one to two years. Once the
seed is ready to be dispersed, the bracts of the female cones open to allow the dispersal of seed;
no fruit formation takes place because gymnosperm seeds have no covering.
Angiosperms vs Gymnosperms
Gymnosperm reproduction differs from that of angiosperms in several ways (figure 10).
In angiosperms, the female gametophyte exists in an enclosed structure—the ovule—which is
within the ovary; in gymnosperms, the female gametophyte is present on exposed bracts of the
female cone. Double fertilization is a key event in the lifecycle of angiosperms but is completely
absent in gymnosperms. The male and female gametophyte structures are present on separate
male and female cones in gymnosperms, whereas in angiosperms, they are a part of the flower.
Lastly, wind plays an important role in pollination in gymnosperms because pollen is blown by
the wind to land on the female cones. Although many angiosperms are also wind-pollinated,
animal pollination is more common.
Figure 10. (a) Angiosperms are flowering plants, and include grasses,
herbs, shrubs, and most deciduous trees, while (b) gymnosperms are
conifers. Both produce seeds but have different reproductive strategies.
ANIMAL REPRODUCTION
The reproductive structures of many animals are very similar, even across different
lineages, in a process that begins with two gametes–eggs and sperm–and ends with a zygote,
which is a fertilized egg. In animals ranging from insects to humans, males
produce sperm in testes, and sperm are stored in the epididymis until ejaculation. Sperm are
small, mobile, low-cost cells that occur in high numbers. Females produce
an ovum or egg that matures in the ovary. Eggs are large cells that require a substantial
investment of time and energy to form, are non-mobile, and are rare relative to sperm numbers.
When the eggs are released from the ovary, they travel to the uterine tubes for fertilization (in
animals that reproduce via internal fertilization) or are released in the aqueous environment (in
animals that reproduce via external fertilization).
All sexual reproducers have females with ovaries that produce large eggs, which
subsequently travel down a uterine tube, and males with testes that produce small, plenteous
sperm, stored in an epididymis. Of course, beyond this general anatomy, there are some
interesting differences in different types of animals:
In some invertebrate species, including many insects and some mollusks and
worms, the female has a spermatheca: a specialized sac which stores sperm for
later use, sometimes up to a year. Fertilization can be timed with environmental or
food conditions that are optimal for offspring survival.
Non-mammal vertebrates, such as most birds and reptiles, have a cloaca: a
single body opening which functions in the digestive, excretory, and reproductive
systems. Mating between birds usually involves positioning the cloaca openings
opposite each other for transfer of sperm from male to female. Ducks are a rare
exception, where the males have a penis.
Mammals have separate openings for the systems in the female, and placental
mammals have a uterus for support of developing offspring. The uterus has two
chambers in species that produce large numbers of offspring at a time, while
species that produce one offspring, such as primates, have a single chamber.
When the sperm have developed flagella and are nearly mature, they leave the testicles
and enter the epididymis, where sperm mature. During ejaculation, the sperm leave the
epididymis and enter the vas deferens, which carries the sperm, behind the bladder, and forms
the ejaculatory duct with the duct from the seminal vesicles.
Semen is a mixture of sperm and spermatic duct secretions and fluids from accessory
glands that contribute most of the semen’s volume. The bulk of the semen comes from the
accessory glands associated with the male reproductive system. These are the seminal vesicles,
the prostate gland, and the bulbourethral gland.
The seminal vesicles are a pair of glands that make thick, yellowish, and alkaline
solution. As sperm are only motile in an alkaline environment, a basic pH is
important to reverse the acidity of the vaginal environment. The solution also
contains mucus, fructose (a source of energy for the sperm cells), a coagulating
enzyme, ascorbic acid (vitamin C), and local-acting hormones called
prostaglandins (may help stimulate smooth muscle contractions in the uterus).
The seminal vesicle glands account for 60 percent of the bulk of semen.
The prostate gland surrounds the urethra, the connection to the urinary bladder.
It has a series of short ducts that directly connect to the urethra. The gland is a
mixture of smooth muscle and glandular tissue. The muscle provides much of the
force needed for ejaculation to occur. The glandular tissue makes a thin, milky
fluid that contains citrate (stimulates sperm motility), enzymes, and prostate
specific antigen (PSA). PSA is a proteolytic enzyme that helps to liquefy the
ejaculate several minutes after release from the male. Prostate gland secretions
account for about 30 percent of the bulk of semen.
The bulbourethral gland releases its secretion prior to the release of the bulk of
the semen. The mucous secretions of this gland help lubricate and neutralize any
acid residue in the urethra left over from urine. This usually accounts for a couple
of drops of fluid in the total ejaculate and may contain a few sperm. Withdrawal
of the penis from the vagina before ejaculation to prevent pregnancy may not
work if sperm are present in the bulbourethral gland secretions.
The table below summarizes the major organs, locations, and functions of male reproductive
anatomy:
Organ Location Function
Scrotum External Carry and supports testes.
Penis External Deliver urine, copulating
organ.
Testes External Produce sperm and male
hormones.
Seminal Vesicles Internal Contribute to semen
production.
Prostate Gland Internal Contribute to semen
production.
Bulbourethral Gland Internal Clean urethra at ejaculation.