Studies in the Alismataceae. XII. Floral organogenesis in Damasonium alisma and Baldellia ranunculoides, and comparisons with Butomus umbellatus

Floral organogenesis of Damasonium alisma Mill. occurs at first in an alternating trimerous pattern typical of Alismataceae, with the formation of three sepals, then three bulges, corresponding to the CA (common perianth–androecium) primordia described in some other Alismataceae, alternating with the sepals. A petal is initiated on each bulge, and a pair of stamens is initiated either on or close to it. Three carpels are initiated in positions alternating with the petals and stamen pairs, and three further carpels then arise above and between the first three. At maturity and in fruit the carpels lie in a whorled arrangement. Floral development in Baldellia ranunculoides (L.) Parl. is identical up to the initiation of the six stamens. After this, six carpel primordia are formed alternating with the stamens, and further carpel primordia arise alternating with those previously formed. In Butomus, up to the initiation of the first six stamens, the general plan of development resembles that of the two Alismataceae. Three further whorls of organs arise in alternation: a whorl of three stamens arises over the stamen pairs followed by two whorls each of three carpel primordia. It is argued that the trimerous appearance of the whorl of sepals (or outer perianth in Butomus) arises de novo and represents a genuine expression of trimery. However, most of the subsequent features of development in these flowers can be seen as arising from phyllotactic mechanisms that cause new primordia to arise between and above pre-existing ones. Consequently the appearance of trimerous or hexamerous whorls above the first whorl of perianth does not represent a fundamental feature of development. The nature of variations in the positional relationships of inner perianth, stamen, and carpel primordia in various Alismataceae and Butomus strengthen the case that there is a significant developmental association between inner perianth members and associated pairs of stamens, which may be connected with the evolution of the flowers from pseudanthial structures.Key words: Baldellia, Butomus, Damasonium, Alismatidae, flower, organogenesis.

Studies in the Alismataceae. X. Floral organogenesis in Luronium natans (L.) Raf.

Floral organogenesis of Luronium natans (L.) Raf. occurs at first in an alternating trimerous pattern typical of Alismataceae, with the formation of three sepals, then three bulges, corresponding to the petal-stamen primordia described in some other Alismataceae, alternating with the sepals. A petal is initiated on each bulge and a pair of stamens is initiated either on it or close to it. After this, development no longer follows a trimerous plan. Six carpels are initiated in positions alternating with the six stamens, and further carpels may then arise above and between the first six. The carpels ultimately lie in a whorled arrangement if there are only six; if more, they may appear whorled or irregularly arranged. After the initiation of the stamen pairs, floral organ primordia appear simply to be positioned between pre-existing primordia as in other phyllotactic systems. It is suggested that the number of carpel primordia formed is probably determined by the size of primordia relative to the floral apex, and the extent of continued growth of the floral apex. Luronium reinforces the concept that a form of trimery is fundamental for the Alismataceae up to the formation of three stamen pairs and adds to the possibilities for variation after this point. It is suggested for the Alismataceae in general that, according to taxon, trimerous development may be terminated at any point after the initiation of the stamen pairs, and after this the primordia are positioned individually in relation to pre-existing primordia. The switch from stamen to carpel initiation is not necessarily correlated with these phyllotactic changes.

Unicarpellate floral development in Potamogeton zosteriformis

Floral development in the species Potamogeton zosteriformis is compared with that of other species of Potamogeton previously investigated (P. densus and P. richardsonii). A spike inflorescence is found on which 6 to 20 flowers develop. These flowers develop their appendages acropetally; first, the four tepals are initiated followed by the four stamens opposite the tepals. As in the flower of P. densus, the two lateral stamens are each initiated as two separate primordia. Unlike any other Potamogeton species where four carpels generally arise, alternating with the tepal and stamen pairs, in P. zosteriformis a single carpel consistently develops on the central portion of the remaining floral meristem. Histological preparations did not reveal any vestigial procambial strands differentiating in the gynoecium. This unique floral development is examined from several points of view. Morphogenetic comparisons are made with other species of Potamogeton which develop tetramerous gynoecia. Morphologically the nature of the Potamogeton flower is reexamined and phylogenetically the possibility of P. zosteriformis as a link or transition between the bisexual and unisexual groups within the Najadales is considered.

Floral organogenesis of Limnocharis flava

Besides a trimerous calyx and corolla, the mature flower exhibits a polyandric androecium and an apocarpous gynoecium consisting of a whorl of carpels. Yet the primary pattern of the flower is completely trimerous and tetracyclic. After the inception of three sepals and three petals, three antesepalous primary androecial primordia are initiated each of which forms three stamens (i.e. secondary androecial primordia). Opposite these three groups of three stamen primordia, three groups of three carpels are initiated, possibly on three extremely inconspicuous primary gynoecial primordia. Additional carpel primordia are formed in varying numbers between the original three groups. Even before carpel inception, the three primary androecial primordia merge laterally thus forming an androecial ring. Additional stamen primordia arise on this ring first between the three groups of three stamen primordia and then in centrifugal direction as the androecial ring broadens basally. Eventually four whorls of stamens and two to three whorls of staminodia are formed secondarily on the androecial ring which arose from the primary primordia. Morphogenesis and construction of the flowers of Limnocharis flava differs in two major respects from those of all other taxa of the Alismatales studied thus far: (1) there are no stamen pairs primarily associated with the petals, and (2) the first-formed carpel primordia do not alternate with the stamen primordia of the preceding whorl, thus violating Hofmeister's rule of alternation.