Anatomical study on the developmental process of the swollen internodes of Phryma (Phrymaceae, eudicots).

We discovered that the internodal swellings of Phryma (eudicots) stems were same as the internodal pulvini of Poaceae (monocots) from the viewpoints of internal structures and functions. The stems of eudicots are usually rod-shaped and are composed of nodes, attached by leaves, and internodes. The internodes of some species, belonging to the clade 'asterids' and its sister clade 'Caryophyllales' of eudicots, have swellings, which have negative tropism, at the basal or apical part of each internode. To know the internal features of the swollen internodes, we performed outer morphological and anatomical studies on the swollen internodes of Phryma, eudicots, one of the genera having swollen internodes, from the winter bud stage to the flowering stage. The results revealed the following: (i) the swollen regions of the internodes were composed of less lignified tissues (e.g., endodermis without Casparian strips, and xylem having less lignified xylem fibers); (ii) the internodal less lignified parts were supported by collenchyma; (iii) the endodermis includes amyloplasts, having accumulated starch granules, which would function as statoliths for negative gravitropism. Consequently, we determined that the swollen parts of the Phryma internodes are same as the internodal pulvini of Poaceae of monocots from the viewpoints of internal structures and functions.