Fertilisation

Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, insemination, pollination, fecundation, syngamy and impregnation, is the fusion of gametes to initiate the development of a new individual organism or offspring. This cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation. In Antiquity, Aristotle conceived the formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in a mode called by him as epigenetic. In 1784, Spallanzani established the need of interaction between the female's ovum and male's sperm to form a zygote in frogs. In 1827, von Baer observed a therian mammalian egg for the first time. Oscar Hertwig (1876), in Germany, described the fusion of nuclei of spermatozoa and of ova from sea urchin. The evolution of fertilisation is related to the origin of meiosis, as both are part of sexual reproduction, originated in eukaryotes. There are two conflicting theories on how the couple meiosis–fertilisation arose. One is that it evolved from prokaryotic sex (bacterial recombination) as eukaryotes evolved from prokaryotes. The other is that mitosis originated meiosis. The gametes that participate in fertilisation of plants are the pollen (male), and the egg (female) cell. Various families of plants have differing methods by which the female gametophyte is fertilized. In Bryophyte land plants, fertilisation takes place within the archegonium. In flowering plants a second fertilisation event involves another sperm cell and the central cell which is a second female gamete. In flowering plants there are two sperm from each pollen grain. In seed plants, after pollination, a pollen grain germinates, and a pollen tube grows and penetrates the ovule through a tiny pore called a micropyle.The sperm are transferred from the pollen through the pollen tube to the ovule. Pollen tube growth Unlike animal sperm which is motile, plant sperm is immotile and relies on the pollen tube to carry it to the ovule where the sperm is released. The pollen tube penetrates the stigma and elongates through the extracellular matrix of the style before reaching the ovary. Then near the receptacle, it breaks through the ovule through the micropyle (an opening in the ovule wall) and the pollen tube 'bursts' into the embryo sac, releasing sperm. The growth of the pollen tube has been believed to depend on chemical cues from the pistil, however these mechanisms were poorly understood until 1995. Work done on tobacco plants revealed a family of glycoproteins called TTS proteins that enhanced growth of pollen tubes. Pollen tubes in a sugar free pollen germination medium and a medium with purified TTS proteins both grew. However, in the TTS medium, the tubes grew at a rate 3x that of the sugar-free medium. TTS proteins were also placed on various locations of semi in vevo pollinated pistils, and pollen tubes were observed to immediately extend toward the proteins. Transgenic plants lacking the ability to produce TTS proteins exhibited slower pollen tube growth and reduced fertility.