Signaling at Physical Barriers during Pollen–Pistil Interactions

In angiosperms, double fertilization requires pollen tubes to transport non-motile sperm to distant egg cells housed in a specialized female structure known as the pistil, mediating the ultimate fusion between male and female gametes. During this journey, the pollen tube encounters numerous physical barriers that must be mechanically circumvented, including the penetration of the stigmatic papillae, style, transmitting tract, and synergid cells as well as the ultimate fusion of sperm cells to the egg or central cell. Additionally, the pollen tube must maintain structural integrity in these compact environments, while responding to positional guidance cues that lead the pollen tube to its destination. Here, we discuss the nature of these physical barriers as well as efforts to genetically and cellularly identify the factors that allow pollen tubes to successfully, specifically, and quickly circumnavigate them.

Paving the Way for Fertilization: The Role of the Transmitting Tract

Angiosperm reproduction relies on the precise growth of the pollen tube through different pistil tissues carrying two sperm cells into the ovules’ embryo sac, where they fuse with the egg and the central cell to accomplish double fertilization and ultimately initiate seed development. A network of intrinsic and tightly regulated communication and signaling cascades, which mediate continuous interactions between the pollen tube and the sporophytic and gametophytic female tissues, ensures the fast and meticulous growth of pollen tubes along the pistil, until it reaches the ovule embryo sac. Most of the pollen tube growth occurs in a specialized tissue—the transmitting tract—connecting the stigma, the style, and the ovary. This tissue is composed of highly secretory cells responsible for producing an extensive extracellular matrix. This multifaceted matrix is proposed to support and provide nutrition and adhesion for pollen tube growth and guidance. Insights pertaining to the mechanisms that underlie these processes remain sparse due to the difficulty of accessing and manipulating the female sporophytic tissues enclosed in the pistil. Here, we summarize the current knowledge on this key step of reproduction in flowering plants with special emphasis on the female transmitting tract tissue.

Stigma, pollen tube transmitting tract, and epidermal micromorphology of the style of Sarracenia purpurea (Sarraceniaceae)

Entomophilous flowers of the genus Sarracenia have a unique umbrella-shaped style, which consists of a broadened and flattened umbrella canopy and a thin cylindrical umbrella stalk. Anatomical and micromorphological features of the style of Sarracenia purpurea L. were studied using light microscopy and scanning electron microscopy. This study found that the pollen tube transmitting tracts (PTTTs) start as a semi-solid canal filled with endotrophic conducting tissue, and run from the peripheral to the center of the canopy where the PTTT becomes a hollow canal supported by ectotrophic conducting tissue. The presence of stomata on the epidermis of the canopy and chloroplasts in its ground parenchyma indicate photosynthetic activities. Convex epidermal cells with intense cuticular striations on the canopy that are similar yet different from those on various regions of the sepals and petals indicate that it may provide contrasting visual cues for pollinators. Multicellular secretory glands and trichomes, which may provide olfactory cues and tactical cues respectively, are also found on the canopy. Thus, the stylar umbrella not only serves as a region for pollen grain capture, pollen germination, and pollen tube transmission but may also play an important role during pollinator–flower interactions.

Localization of membrane-associated calcium in unpollinated and pollinated pistil of Petunia hybrida Hort.

In the pistil of <em>Petunia hybrida</em>, the transmitting tract and the ovules are the sites which give Ca²⁺-CTC fluorescence. In unpollinated pistil the level of membrane-associated Ca²⁺ decreases from the stigma to the base of the style. The renewed strong rise of Ca²⁺-CTC fluorescence appears on the placenta surface and in the ovule integuments. Following pollination, when the pollen tubes have grown through the pistil, the pattern of membrane-associated Ca²⁺ on the path stigma - ovary is reversed. The highest fluorescence is found in the base of the style. In pollinated ovules the Ca²⁺-CTC fluorescence increases markedly. In the transmitting cells membrane-associated Ca²⁺ occurs mainly in the polar regions of the cell. During cell degeneration following pollination the cytoplasmic clusters show Ca²⁺-CTC fluorescence. The used <em>P. hybrida</em> cultivar is self-fertile. The selection of pollen tubes occurs mainly in the upper part of the style. The rejected pollen tubes show a steady high level of membrane-associated calcium.