Cytoarchitecture of Utricularia nutritive tissue.

Beginning with light microscopy studies in the late 19th century, the placental "nutritive tissue" in carnivorous plants of Utricularia spp. has been well described by several authors. Based on observations of direct contact between the embryo sac and the "nutritive tissue" and the lack of vascularization of the ovule, it has been suggested that this nutritive tissue plays a key role in the nutrition of the female gametophyte. To date, however, the structure of this tissue has received only scant attention. To fill this knowledge gap, we have characterized its anatomy and histochemistry in more detail and addressed the speculations of a number of earlier researchers. Nutritive tissue during the period of flower opening in three Utricularia species, each belonging to different sections and subgenera (Polypompholyx, Bivalvaria and Utricularia), was examined by light and, in particular, electron microscopy. In all of the investigated species, nutritive tissue cells differ from placental parenchyma cells in having no huge vacuole, no large amyloplasts with starch grains, and no protein inclusions in the nucleus. The funicular nutritive tissue in U. dichotoma consists of active cells with a secretory character, while U. sandersonii has a small placental nutritive tissue consisting of colenchymatous cells accumulating lipids. The most complex nutritive tissue occurs in aquatic U. intermedia, which occupies a derived position in the genus phylogeny. In this latter species, the cells of this tissue resemble meristematic cells in having a relatively large nucleus, thin cell walls, and reduced vacuoles, but the well-developed endoplasmic reticulum (ER) in some cells is similar to that in secretory cells. The cytoplasm is rich in microtubules, some of which are in close contact with the ER cisternae. We found very thick cell walls between nutritive tissue cells and parenchyma cells, but plasmodesmata between these types of cells are rare. Similarities in both the position and structure of nutritive tissue in Polypompholyx and section Pleiochasia support their classification together in one subgenus, based on results from a molecular study. The position and structure of the nutritive tissue in Utricularia spp. are related to the position of various species in the genus phylogeny.