Homeosis has played an important role in the evolution of the flowers of the Zingiberales, especially those of the Ginger Group. In the Zingiberaceae, two members of the outer androecial whorl are replaced by a lip, and two members of the inner androecial whorl are replaced by petaloid staminodes. Most of the androecium of the Costaceae has also been replaced by petaloid structures, and the single fertile stamen is often attached to an enlarged petaloid “filament.” The Cannaceae and Marantaceae have one‐half of one fertile anther and three to four variously modified staminodes. In contrast, homeosis has played a minor role in floral evolution of the Banana Group. Only in the Heliconiaceae has a stamen been replaced by a staminode. In none of the families of the Zingiberales do the staminodes assume the total “form or character” of any perianth members. Because of this, it is reasonable to extend the definition of homeosis to include replacement by an organ like, but not identical to, some other part of the plant.
Hofmeister's rule is an empirical heuristic derived from the observation that new leaf primordia are formed in the largest space between the existing flanks of the older primordia. These observations have been repeatedly validated in studies of leaf arrangement, but there has been little attempt to extend them to inflorescence and floral organs. This investigation demonstrates the validity of Hofmeister's observations to cincinnus and early flower development in Phenakospermum guyannense (Strelitziaceae) and Heliconia latispatha (Heliconiaceae) and relates these results to Paul Green's work on the biophysics of organ formation. The cincinni of Phenakospermum and Heliconia arise in the axils of primary bracts and produce a prophyll, continuation apex, and flower in regular succession. The shapes and orientations of the apical regions of the cincinni are correlated with the placement of these organs, which in turn effect the positions of the sepals and their sequence of formation. The result is two rows of mirror-image flowers. The mirror-image symmetry of the flowers is a direct result of Hofmeister's rule in connection with the shape of the apical region. These two factors create a self-sustaining developmental system that produces prophylls, continuation apices, flowers, and sepals in regular succession. Although the cincinni and flowers of these species are built on a common plan, slight differences in apical shape and orientation produce differences in mature floral orientation. Understanding these orientations, and identifying the sequence of sepal formation, allows a proper identification of organ homologies. A study of Green's results and theories shows that Hofmeister's rule can be considered as an empirical condensation of the biophysical factors that influence organ position. These biophysical factors are widely applicable to organ formation in many species.
Flower organogenesis in Hedychium gardnerianum and Hedychium coronarium begins with the sequential formation of the sepals on a rounded–triangular apex. Growth in three regions of the apex, inside the sepals, produces three common petal – inner androecial primordia. Intercalary growth below and between these primordia produces a floral cup, the site of gynoecial formation. The common primordia separate to form petals and inner androecial members. After separation, the anterior inner androecial members fuse to form the labellum. The posterior member forms the polleniferous stamen. Up to this point in development, the flowers of H. coronarium are slightly ahead of those of H. gardnerianum. However, in H. gardnerianum, initiation of the two thecae of the stamen occurs immediately following the separation of the common primordia, while thecae formation in H. coronarium is slightly delayed. These results show that the relative timing of developmental events can vary even over a short developmental period. Formation of the outer androecium takes place at the apices of a triangle formed by the inner androecial members. The anterior outer androecial member aborts soon after formation, whereas the posterior outer androecial members form the two petaloid staminodes that are found on the lateral sides of the labellum in the mature flower. Key words: flower development, flower structure, inflorescence, stamen, Zingiberaceae, Hedychium.
This paper discusses problems with labelling plant structures in the context of attempts to create a unified Plant Structure Ontology. Special attention is given to structures with mixed, or doubtful identities that are difficult or even impossible to label with a single term. In various vascular plants (and some groups of animals) the structural categories for the description of forms are less distinct than is often supposed. Thus, there are morphological misfits that do not fit exactly into one or the other category and to which it is difficult, or even impossible, to apply a categorical name. After presenting three case studies of intermediate organs and organs whose identity is in doubt, we review five approaches to categorizing plant organs, and evaluate the potential of each to serve as a general reference system for gene annotations. The five approaches are (1) standardized vocabularies, (2) labels based on developmental genetics, (3) continuum morphology, (4) process morphology, (5) character cladograms. While all of these approaches have important domains of applicability, we conclude that process morphology is the one most suited to gene annotation.
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