13.1 Reproductive development and structure  (Page 3/30)

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 ( [link] ). 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    ( [link] ). 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 ( [link] ). 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.