169 Reproductive Development and Structure

Learning Objectives

By the end of this section, you will be able to do the following:

  • Describe the two stages of a plant’s lifecycle
  • Compare and contrast male and female gametophytes and explain how they form in angiosperms
  • Describe the reproductive structures of a plant
  • Describe the components of a complete flower
  • Describe the development of microsporangium and megasporangium in gymnosperms

Sexual reproduction takes place with slight variations in different groups of plants. Plants have two distinct stages in their lifecycle: the gametophyte stage and the sporophyte stage. The haploid gametophyte produces the male and female gametes by mitosis in distinct multicellular structures. Fusion of the male and females gametes forms the diploid zygote, which develops into the sporophyte. After reaching maturity, the diploid sporophyte produces spores by meiosis, which in turn divide by mitosis to produce the haploid gametophyte. The new gametophyte produces gametes, and the cycle continues. This is the alternation of generations, and is typical of plant reproduction ((Figure)).

The alternation of generations in angiosperms is depicted in this diagram. (credit: modification of work by Peter Coxhead)


Illustration shows the life cycle of angiosperms, which includes a microgametophyte stage and a megagametophyte stage. The life cycle begins with the fusion of egg and sperm to form a zygote. The zygote undergoes mitosis, resulting in a male microsporophyte or a female megasporophyte. The microsporophyte has a cluster of cells called a microsporangium, and the megasporophyte has a cluster of cells called a megasporangium. Through meiosis, the microsporangium forms microspores, and the megasporangium forms megaspores. Both microspores and megaspores undergo mitosis, forming the microgametophyte and megagametophyte, respectively. Within the microgametophyte, the fusion of egg and sperm completes the cycle.

The life cycle of higher plants is dominated by the sporophyte stage, with the gametophyte borne on the sporophyte. In ferns, the gametophyte is free-living and very distinct in structure from the diploid sporophyte. In bryophytes, such as mosses, the haploid gametophyte is more developed than the sporophyte.

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.

Sexual Reproduction in Angiosperms

The lifecycle of angiosperms follows the alternation of generations explained previously. The haploid gametophyte alternates with the diploid sporophyte during the sexual reproduction process of angiosperms. Flowers contain the plant’s reproductive structures.

Flower Structure

A typical flower has four main parts—or whorls—known as the calyx, corolla, androecium, and gynoecium ((Figure)). The outermost whorl of the flower has green, leafy structures known as sepals. The sepals, collectively called the calyx, help to protect the unopened bud. The second whorl is comprised of petals—usually, brightly colored—collectively called the corolla. The number of sepals and petals varies depending on whether the plant is a monocot or dicot. In monocots, petals usually number three or multiples of three; in dicots, the number of petals is four or five, or multiples of four and five. Together, the calyx and corolla are known as the perianth. The third whorl contains the male reproductive structures and is known as the androecium. The androecium has stamens with anthers that contain the microsporangia. The innermost group of structures in the flower is the gynoecium, or the female reproductive component(s). The carpel is the individual unit of the gynoecium and has a stigma, style, and ovary. A flower may have one or multiple carpels.

Visual Connection
The four main parts of the flower are the calyx, corolla, androecium, and gynoecium. The androecium is the sum of all the male reproductive organs, and the gynoecium is the sum of the female reproductive organs. (credit: modification of work by Mariana Ruiz Villareal)


Illustration shows parts of a flower, which is called the perianth. The corolla is composed of petals, and the calyx is composed of sepals. At the center of the perianth is a vase-like structure called the carpel. A flower may have one or more carpels, but the example shown has only one. The narrow neck of the carpel, called the style, widens into a flat stigma at the top. The ovary is the wide part of the carpel. Ovules, or megasporangia, are clusters of pods in the middle of the ovary. The androecium is composed of stamens which cluster around the carpel. The stamen consists a long, stalk-like filament with an anther at the end. The anther shown is tri-lobed. Each lobe, called a microsporangium, is filled with pollen.

If the anther is missing, what type of reproductive structure will the flower be unable to produce? What term is used to describe an incomplete flower lacking the androecium? What term describes an incomplete flower lacking a gynoecium?

<!–<solution> Pollen (or sperm); carpellate; staminate.
–>

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)).

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 flowers are clustered in the immature ears. Each strand of silk is a stigma. The corn kernels are seeds that develop on the ear after fertilization. Also shown is the lower stem and root.


Illustration shows parts of a corn plant. Staminate flowers are tiny flowers that cluster in strands to form the tassel at the top of the plant. Pollen grains are small, teardrop-shaped structures. Carpellate flowers are clustered in the immature ear, which is covered by leaves. Silk protrudes from the tops of the leaves covering the flower. In the mature ear, the kernels form where the carpellate flowers were located.

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)).

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. (credit a photo: modification of work by Benjamin Zwittnig; credit b photo: modification of work by “Koshy Koshy”/Flickr)


Part A shows a lily, which has an ovary above the petals. The ovary sits above the teardrop-shaped petals. Part B shows several fuchsia flowers hanging down from a stem. The ovary is below the edge of the petals.

Male Gametophyte (The Pollen Grain)

The male gametophyte develops and reaches maturity in an immature anther. In a plant’s male reproductive organs, development of pollen takes place in a structure known as the microsporangium ((Figure)). The microsporangia, which are usually bilobed, are pollen sacs in which the microspores develop into pollen grains. These are found in the anther, which is at the end of the stamen—the long filament that supports the anther.

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, as shown in the (b) micrograph of an immature lily anther. In these scanning electron micrographs, pollen sacs are ready to burst, releasing their grains. (credit a: modification of work by LibreTexts; b: modification of work by Robert R. Wise; scale-bar data from Matt Russell)


Illustration A shows cross section of an anther, which has four lobes each containing a pollen sac, or microsporangium. Inside the pollen sac is a layer called the tapetum, and within this ring are the microspore mother cells. As the microsporangium matures, two pollen sacs merge and an opening forms between them so that the pollen can be released. Micrographs in part B show pollen sacs with a visible opening between them. b: A micrograph of an immature lily anther shows four pollen sacs containing pollen 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)). 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.

Pollen develops from the microspore mother cells. The mature pollen grain is composed of two cells: the pollen tube cell and the generative cell, which is inside the tube cell. The pollen grain has two coverings: an inner layer (intine) and an outer layer (exine). The inset scanning electron micrograph shows Arabidopsis lyrata pollen grains. (credit “pollen micrograph”: modification of work by Robert R. Wise; scale-bar data from Matt Russell)


Illustration shows the formation of pollen from a microspore mother cell. The mother cell undergoes meiosis to form a tetrad of cells, which separate to form the pollen grains. The pollen grains undergo mitosis without cytokinesis, resulting in four mature pollen grains with two nuclei each. One is called the generative nucleus, and the other is called the pollen tube nucleus. Two projective layers form around the mature pollen grain, the inner intine and the outer exine. Micrograph shows a pollen grain, which looks like puffed wheat.

Each pollen grain has two coverings: the exine (thicker, outer layer) and the intine ((Figure)). 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)

While the details may vary between species, the overall development of the female gametophyte has two distinct phases. First, in the process of megasporogenesis, a single cell in the diploid megasporangium—an area of tissue in the ovules—undergoes meiosis to produce four megaspores, only one of which survives. During the second phase, megagametogenesis, the surviving haploid megaspore undergoes mitosis to produce an eight-nucleate, seven-cell female gametophyte, also known as the megagametophyte or embryo sac. Two of the nuclei—the polar nuclei—move to the equator and fuse, forming a single, diploid central cell. This central cell later fuses with a sperm to form the triploid endosperm. Three nuclei position themselves on the end of the embryo sac opposite the micropyle and develop into the antipodal cells, which later degenerate. The nucleus closest to the micropyle becomes the female gamete, or egg cell, and the two adjacent nuclei develop into synergid cells ((Figure)). The synergids help guide the pollen tube for successful fertilization, after which they disintegrate. Once fertilization is complete, the resulting diploid zygote develops into the embryo, and the fertilized ovule forms the other tissues of the seed.

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.

Visual Connection
As shown in this diagram of the embryo sac in angiosperms, the ovule is covered by integuments and has an opening called a micropyle. Inside the embryo sac are three antipodal cells, two synergids, a central cell, and the egg cell.


Illustration depicts the embryo sac of an angiosperm, which is egg-shaped. The narrow end, called the micropylar end, has an opening that allows pollen to enter. The other end is called the chalazal end. Three cells called antipodals are at the chalazal end. The egg cell and two other cells called synergids are at the micropylar end. Two polar nuclei are inside the central cell in the middle of the embryo sac.

An embryo sac is missing the synergids. What specific impact would you expect this to have on fertilization?

  1. The pollen tube will be unable to form.
  2. The pollen tube will form but will not be guided toward the egg.
  3. Fertilization will not occur because the synergid is the egg.
  4. Fertilization will occur but the embryo will not be able to grow.

<!– <para>B–>

Sexual Reproduction in Gymnosperms

As with angiosperms, the lifecycle of a gymnosperm is also characterized by alternation of generations. In conifers such as pines, the green leafy part of the plant is the sporophyte, and the cones contain the male and female gametophytes ((Figure)). The female cones are larger than the male cones and are positioned towards the top of the tree; the small, male cones are located in the lower region of the tree. Because the pollen is shed and blown by the wind, this arrangement makes it difficult for a gymnosperm to self-pollinate.

This image shows the lifecycle 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. (credit “female”: modification of work by “Geographer”/Wikimedia Commons; credit “male”: modification of work by Roger Griffith)


The conifer life cycle begins with a mature tree, which is called a sporophyte and is diploid, 2 n. The tree produces male cones in the lower branches, and female cones in the upper branches. The male cones produce pollen grains that contain two generative, sperm, nuclei and a tube nucleus. When the pollen lands on a female scale, a pollen tube grows toward the female gametophyte, which consists of an ovule containing the megaspore. Upon fertilization, a diploid zygote forms. The resulting seeds are dispersed, and grow into a mature tree, ending the cycle. Both the male and female cone are made up of rows of scales, but the female cone is round and wide, and the male cone is long and thin with thinner scales.

Male Gametophyte

A male cone has a central axis on which bracts, a type of modified leaf, are attached. The bracts are known as microsporophylls ((Figure)) 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.

Link to Learning

Watch this video to see a cedar releasing its pollen in the wind.

Female Gametophyte

The female cone also has a central axis on which bracts known as megasporophylls ((Figure)) 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.

This series of micrographs shows male and female gymnosperm gametophytes. (a) This male cone, shown in cross section, has approximately 20 microsporophylls, 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. (credit: modification of work by Robert R. Wise; scale-bar data from Matt Russell)


Part a shows a cross section of a male cone, which is oval with a flattened bottom. A stem-like structure runs up the middle, and oblong microsporophylls radiate from either side. Migrograph b shows a microsphorphyll, which is filled with round pollen grains. Micrograph C shows a pollen grain, which is oval with two lobes attached. Part D shows a cross section of a female cone, which is similar in shape to the male cone but with a wider central structure. Megasporophylls radiate from either side of this structure. At the base the megasprophylls are narrow, and the widen out into a roughly diamond shape. Part E shows the megasprophyll, which has an oval ovule at its base. Part F shows the ovule. The megaspore mother cell is at the center. An opening called a micropyle allows entry of the pollen tube from the base.

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 versus Gymnosperms

Gymnosperm reproduction differs from that of angiosperms in several ways ((Figure)). 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.

(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. (credit a: modification of work by Wendy Cutler; credit b: modification of work by Lews Castle UHI)


Photo A shows a deciduous tree that loses its leaves in winter. Photo B shows a conifer: a tree that has needles year round.

Link to Learning

View an animation of the double fertilization process of angiosperms.

Section Summary

The flower contains the reproductive structures of a plant. All complete flowers contain four whorls: the calyx, corolla, androecium, and gynoecium. The stamens are made up of anthers, in which pollen grains are produced, and a supportive strand called the filament. The pollen contains two cells— a generative cell and a tube cell—and is covered by two layers called the intine and the exine. The carpels, which are the female reproductive structures, consist of the stigma, style, and ovary. The female gametophyte is formed from mitotic divisions of the megaspore, forming an eight-nuclei ovule sac. This is covered by a layer known as the integument. The integument contains an opening called the micropyle, through which the pollen tube enters the embryo sac.

The diploid sporophyte of angiosperms and gymnosperms is the conspicuous and long-lived stage of the life cycle. The sporophytes differentiate specialized reproductive structures called sporangia, which are dedicated to the production of spores. The microsporangium contains microspore mother cells, which divide by meiosis to produce haploid microspores. The microspores develop into male gametophytes that are released as pollen. The megasporangium contains megaspore mother cells, which divide by meiosis to produce haploid megaspores. A megaspore develops into a female gametophyte containing a haploid egg. A new diploid sporophyte is formed when a male gamete from a pollen grain enters the ovule sac and fertilizes this egg.

Visual Connection Questions

(Figure) If the anther is missing, what type of reproductive structure will the flower be unable to produce? What term is used to describe a flower that is normally lacking the androecium? What term describes a flower lacking a gynoecium?

(Figure) Pollen (or sperm); carpellate; staminate.

(Figure) An embryo sac is missing the synergids. What specific impact would you expect this to have on fertilization?

  1. The pollen tube will be unable to form.
  2. The pollen tube will form but will not be guided toward the egg.
  3. Fertilization will not occur because the synergid is the egg.
  4. Fertilization will occur but the embryo will not be able to grow.

(Figure) B: The pollen tube will form but will not be guided toward the egg.

Review Questions

In a plant’s male reproductive organs, development of pollen takes place in a structure known as the ________.

  1. stamen
  2. microsporangium
  3. anther
  4. tapetum

B

The stamen consists of a long stalk called the filament that supports the ________.

  1. stigma
  2. sepal
  3. style
  4. anther

D

The ________ are collectively called the calyx.

  1. sepals
  2. petals
  3. tepals
  4. stamens

A

The pollen lands on which part of the flower?

  1. stigma
  2. style
  3. ovule
  4. integument

A

Critical Thinking Questions

Describe the reproductive organs inside a flower.

Inside the flower are the reproductive organs of the plant. The stamen is the male reproductive organ. Pollen is produced in the stamen. The carpel is the female reproductive organ. The ovary is the swollen base of the carpel where ovules are found. Not all flowers have every one of the four parts.

Describe the two-stage lifecycle of plants: the gametophyte stage and the sporophyte stage.

Plants have two distinct phases in their lifecycle: the gametophyte stage and the sporophyte stage. In the gametophyte stage, when reproductive cells undergo meiosis and produce haploid cells called spores, the gametophyte stage begins. Spores divide by cell division to form plant structures of an entirely new plant. The cells in these structures or plants are haploid. Some of these cells undergo cell division and form sex cells. Fertilization, the joining of haploid sex cells, begins the sporophyte stage. Cells formed in this stage have the diploid number of chromosomes. Meiosis in some of these cells forms spores, and the cycle begins again: a process known as alternation of generations.

Describe the four main parts, or whorls, of a flower.

A typical flower has four main parts, or whorls: the calyx, corolla, androecium, and gynoecium. The outermost whorl of the flower has green, leafy structures known as sepals, which are collectively called the calyx. It helps to protect the unopened bud. The second whorl is made up of brightly colored petals that are known collectively as the corolla. The third whorl is the male reproductive structure known as the androecium. The androecium has stamens, which have anthers on a stalk or filament. Pollen grains are borne on the anthers. The gynoecium is the female reproductive structure. The carpel is the individual structure of the gynoecium and has a stigma, the stalk or style, and the ovary.

Discuss the differences between a complete flower and an incomplete flower.

If all four whorls of a flower are present, it is a complete flower. If any of the four parts is missing, it is known as incomplete. Flowers that contain both an androecium and gynoecium are called androgynous or hermaphrodites. Those that contain only an androecium are known as staminate flowers, and those that have only carpels are known as carpellate. If both male and female flowers are borne on the same plant, it is called monoecious, while plants with male and female flowers on separate plants are termed dioecious.

Glossary

androecium
sum of all the stamens in a flower
antipodals
the three cells away from the micropyle
exine
outermost covering of pollen
gametophyte
multicellular stage of the plant that gives rise to haploid gametes or spores
gynoecium
the sum of all the carpels in a flower
intine
inner lining of the pollen
megagametogenesis
second phase of female gametophyte development, during which the surviving haploid megaspore undergoes mitosis to produce an eight-nucleate, seven-cell female gametophyte, also known as the megagametophyte or embryo sac.
megasporangium
tissue found in the ovary that gives rise to the female gamete or egg
megasporogenesis
first phase of female gametophyte development, during which a single cell in the diploid megasporangium undergoes meiosis to produce four megaspores, only one of which survives
megasporophyll
bract (a type of modified leaf) on the central axis of a female gametophyte
micropyle
opening on the ovule sac through which the pollen tube can gain entry
microsporangium
tissue that gives rise to the microspores or the pollen grain
microsporophyll
central axis of a male cone on which bracts (a type of modified leaf) are attached
perianth
(also, petal or sepal) part of the flower consisting of the calyx and/or corolla; forms the outer envelope of the flower
polar nuclei
found in the ovule sac; fusion with one sperm cell forms the endosperm
sporophyte
multicellular diploid stage in plants that is formed after the fusion of male and female gametes
synergid
type of cell found in the ovule sac that secretes chemicals to guide the pollen tube towards the egg

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