thumb|SEM image of pollen tubes growing from lily pollen grains.

A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells from the pollen grain—either from the stigma (in flowering plants) to the ovules at the base of the pistil or directly through ovule tissue in some gymnosperms. In maize, this single cell can grow longer than to traverse the length of the pistil.

Pollen tubes were first discovered by Giovanni Battista Amici in the 19th century.

They are used as a model for understanding plant cell behavior. Research is ongoing to comprehend how the pollen tube responds to extracellular guidance signals to achieve fertilization.

thumb|Bee pollinating a sunflower. Pollen is transferred from anther of one plant to stigma of another as bee collects nectar

Pollen tubes are unique to seed plants and their structures have evolved over their history since the Carboniferous period. Pollen tube formation is complex and the mechanism is not fully understood.

The germinated pollen tube must drill its way through the nutrient-rich style and curl to the bottom of the ovary to reach an ovule. Once the pollen tube reaches an ovule, it bursts to deliver the two sperm cells. One of the sperm cells fertilizes the egg cell which develops into an embryo, which will become the future plant. The other one fuses with both polar nuclei of the central cell to form the endosperm, which serves as the embryo's food supply. Finally, the ovary will develop into a fruit and the ovules will develop into seeds.

Gymnosperms

thumb|186x186px|Cross section of ovule in gymnosperms and angiosperms

Gymnosperm pollen is produced in microsporangia borne on the scales of the male cone or microstrobilus. In most species, the plants are wind-pollinated, and the pollen grains of conifers have air bladders that provide buoyancy in air currents. The grains are deposited in the micropyle of the ovule of a female cone or megastrobilus, where they mature for up to a year. In conifers and gnetophytes, the pollen germinate to produce a pollen tube that penetrates the megasporangium or nucellus carrying with it sperm nuclei that are transferred to the egg cell in the developing archegonia of the female plant. The apex region is where tip growth occurs and requires the fusion of secretory vesicles. There is mostly pectin and homogalacturonans (part of the cell wall at the pollen tube tip) inside these vesicles. The pectin in the apex region contains methylesters which allow for flexibility, before the enzyme pectin methylesterase removes the methylester groups allowing calcium to bind between pectins and give structural support. The homogalacturonans accumulate in the apex region via exocytosis in order to loosen the cell wall. A thicker and softer tip wall with a lower stress yield will form and this allows cell expansion to occur, which leads to an increase in tip growth. Reverse-fountain cytoplasmic streaming occurs during the tip growth which is essential for the cellular expansion, because it is transporting organelles and vesicles between the shank region and subapex region.

The actin cytoskeleton is an important factor in pollen tube growth, because there are different patterns of actin cytoskeleton within the different regions of the pollen tube for the maintenance of polarized cell growth. For instance, there are longitudinal actin cables in the shank region in order to regulate reverse-fountain cytoplasmic streaming. The F-actin controls the accumulation of the homogalacturonans full vesicles- essentially mediating tip growth- in the subapex region. The actin filaments controls the apical membrane and cytoplasm interactions while the pollen tube is growing in the apex region. The F-actin from the apical membrane makes an actin binding protein called formin which is essential for pollen tube tip growth. Formins are expressed in the tip growth cells and are divided into two subgroups: type I and type II. The type I formins make the actin structures and partake in cytokinesis. The type II formins on the other hand contribute to the growth of polarized cells which is necessary for tip growth. Tip growth is a form of extreme polarized growth and this polarized process requires actin-binding protein-mediated organization of actin cytoskeleton. An essential protein required for this tip growth is the actin-organizing protein and type II formin protein called Rice Morphology Determinant (RMD). RMD is localized in the tip of the pollen tube and controls pollen tube growth by regulating the polarity and organization of F-actin array.

RMD promotes pollen tube growth

RMD promotes pollen germination and pollen tube growth, and this is proven through numerous experiments. The first experiment compares the features of the pistil and the stigma of rmd-1 mutant (rice plant without a functional RMD) and the wild-type rice plant (with a functional RMD). The anther and pistil were shorter in the rmd-1 mutants than the wild-type. This experiment showed that RMD is critical for pollen development. Wild-type rice plants have increased germination rates while rmd-1 mutants have decreased germination rates. This was seen when both were germinated in a liquid germination medium. After the germination rates were tested, there was a comparison of the lengths and widths of the pollen tubes between the two plants. The pollen tubes of the wild-type plants had a greater pollen tube length than the mutants, but the mutants had a greater tube width. This greater pollen tube width within the mutants indicates the decrease in the growth of polarized cells and thus decrease in tip growth. Next, pollen grains from the wild type and mutants were collected to compare the pollination activities between the wild types and mutants. There was decreased activity and minimal penetration within the mutants whereas an increased activity and penetration through the style and to the bottom of the pistils within the wild types. These observations indicated the delayed pollen tube growth in the rmd-1 mutants. Additionally, there was no effect on fertilization rates between the wild type and the mutant and this was tested by measuring the seed-setting rates between the wild type and mutant. It was found that both had similar seed-setting rates. Therefore, RMD does not affect fertilization and has an effect only on tip growth. There was greater fluorescence intensity in the shank region of the rmd-mutant tubes which means there was a higher density of F-actin within this region. But, there was a lower density of F-actin observed in the tip region of the rmd-mutant tubes compared to the wild type tubes. This demonstrates that the F-actin distribution pattern of pollen tubes is altered without a functional RMD.

In order to determine the polarity of the actin cables, the angles between the actin cables and elongation axis of the pollen tube were measured. The angles in the shank region of the wild type pollen tubes were predominantly less than 20° whereas the angles for the rmd-mutant pollen tubes were greater than 60°. These results support the fact that RMD is essential for polarized tip growth, because the rmd-mutant pollen tubes (without a functional RMD) exhibited an increased width, and thus a decrease in tip growth. The maximum length of the single cables of F-actin filaments from the apical to the shank region of elongating pollen tubes were also measured to test the polarity within the pollen tube. The maximum length of the F-actin cables were shorter in the rmd-mutant pollen tubes compared to those in the wild type tubes. Therefore, these combined results support that the proper organization of actin cables as well as normal F-actin densities within the tip of the tube can only be achieved if RMD is present.

See also

  • Evolutionary history of plants
  • Flowering plant
  • Self-incompatibility in plants
  • Siphonogamy

References

  • Pollen tube primer
  • Images : Pollen tetrad and Pollen tube Calanthe discolor Lindl. - Flavon's Secret Flower Garden