Ryoko Imaichi

Comparative leaf development ofOsmunda lancea andO. japonica (osmundaceae): Heterochronic origin of rheophytic stenophylly

Comparative development of the narrow pinnules of rheophyticOsmunda lancea and of the broad pinnules of a related dryland species,O. japonica, was examined and the origin of rheophytic stenophylly was discussed. The mature leaves and their various parts ofO. lancea are smaller and narrower than those ofO. japonica. The young pinnules ofO. lancea at the initiation of cell expansion are smaller than those ofO. japonica. The growth pattern of the pinnules is fundamentally the same in the two species, but pinnule growth period is shorter inO. lancea than inO. japonica. While the largest growth rate in pinnule length is quite similar, inO. lancea the pinnules are less elongated and much less broadened during ontogeny. Cell expansion in the mesophyll and epidermis proceeds acropetally and toward the margin along the axes of costules and veins. Although the numbers of mesophyll and epidermal cells between two adjacent veinlets are almost the same inO. lancea andO. japonica, during the subsequent growth period inO. lancea, the cells expand to a smaller extent and the veinlets become more narrowly oblique to the costule. This oblique distortion of laminar segments framed by veins causes stenophylly, an allometric modification. The stenophylly ofO. lancea is believed to have arisen by heterochronic evolution, in particular, progenesis.

Leaf morphology of a facultative rheophyte,Farfugium japonicum var.luchuense (Compositae)

The gross morphology and anatomy of leaves in a facultative rheophyte,Farfugium japonicum var.luchuense, were examined in order to clarify how the two characters are correlated with one another and how the facultative rheophyte differs from obligate rheophytes in leaf morphology and anatomy. Most rheophytes of the variety have narrowly cuneate leaf base, while most inland plants usually have truncate to cordate one, although the habitat-morphology correlation is not so clear. The leaf shape or the divergence angle of leaf base is strongly correlated with the number of primary veins and intervein-distance dependent on the number of cells intervening between veins. This makes a marked contrast to many reported cases of obligate rheophytes in which the leaf shape is strongly correlated with cell size. There is a rough tendency that narrower leaves of rheophytes have a thicker cuticular layer. However, the cell size and the volume (area) of intercellular space differ only slightly with the leaf shape.

Developmental anatomy of the shoot apical cell, rhizophore and root ofSelaginella uncinata

The developmental anatomy of the shoot apex, rhizophore and root ofSelaginella uncinata was examined by the semi-thin section method. The shoot apex has a single, lens-shaped apical cell with two cutting faces. Rhizophore primordia are initiated exogenously at the branching point of the second youngest lateral shoot. The rhizophore apex has a tetrahedral apical cell with three cutting faces. A pair of root primordia is initiated endogenously from inner cells of the rhizophore apex, after the rhizophore apical cell becomes unidentifiable losing its activity, and subsequently a root cap is formed from the distal face of the root apical cell. During the course of successive root branching the apical cell in an original root apical meristem becomes unidentifiable and then a new apical cell is initiated in each of the bifurcated root apical meristems. The root branching mode seems to be equivalent to the described dichotomous branching mode of fern shoots. Our results demonstrate a distinct morphogenetical difference between the rhizophore and the root, and confirm the exogenous origin of the rhizophore, as described for other species ofSelaginella. This evidence indicates that the rhizophore is not an aerial root but a leafless, root-producing axial organ.

Ovular Development and Morphology in Some Magnoliaceae Species

Floral phenology and ovular development ofLiriodendron tulipifera are described. The ovule primordia are initiated in December, followed by prominent development in March, and the ovules are mature in May. The inner integument is formed as an annular rim on the incurving ovule primordia, but the outer integument develops as a semi-annular rim interrupted on the concave side of the funicle. Later, an outgrowth, which is interpreted here as an obturator, arises on the concave side of the funicle. The funicular outgrowth arises far from the inner integument, while the outer integument is close to the inner. The outer integument and the funicular outgrowth together form an envelope complex. Later the outer integument produces two distal lobes, which disappear at maturity. Mature ovules of the threeMagnolia species examined have similar lobes. It is suggested that the hood-shaped outer integument is primitive in angiosperms.

Developmental studies on the leaf and the extra-axillary bud ofHistiopteris incisa

Anatomical and developmental studies have been made ofHistiopteris incisa in order to obtain a reasonable interpretation of the so-called extra-axillary bud. Single, or rarely two extra-axillary buds arise on the lateral side of the petiolar base. The branch trace appears to depart from the basiscopic margin of the leaf trace. At the earliest stage of the leaf initiation, the leaf apical cell is cut off in one of the prismatic cells of the shoot apical meristem. The leaf apical cell, then, cuts off segments successively to form a well-defined group of derivatives. On the other hand, a well-recognized cell group called “outer neighboring cell group”,onc, is found adjacent to the abaxial boundary of the derivatives of the leaf apical cell. This group of cells does not originate directly in the mother cell of the leaf apical cell. The primordium of the extra-axillary bud is always initiated in the superficial pillar-shaped cell layer ofonc. The leaf primordium may consist of two parts, the distal part derived from the leaf apical cell and the basal part from the adjacent cells includingonc. These facts suggest that the extra-axillary bud is of foliar nature.

Comparative leaf morphology of young sporophytes of rheophyticOsmunda lancea and drylandO. japonica

Field and morphological observations were made of the young sporophytes of rheophyticOsmunda lancea and its related drylandO. japonica, and the rheophytes adaptation in the early sporophytic stages was discussed. Mature plants ofO. lancea andO. japonica do not occur in dryland and rheophytic habitats, respectively, but their very young sporophytes rarely grow there. The young sporophytes ofO. lancea differ considerably from those ofO. japonica in having the relatively short petioles with thin-walled epidermal cells, early lamina partition, cuneate leaf- and pinna-base, oblique (not horizontal) lamina disposition, a fine network of spongy tissue in the 4th and older leaves, and dense epicuticular wax deposits on leaf epidermis. They seem to relate to the flexibility of petioles and the toughness and flood-tolerance of blades, and make the young sporophytes adapted to the rheophytic habitat.Osmunda japonica lacking those characteristics disappears from the rheophytic habitat during the early ontogenetic stages.

Developmental anatomy of the three-dimensional leaf of Botrychium ternatum (Thunb.) Sw.

The ophioglossaceous leaf exhibits a unique morphology, three-dimensional constitution. This examination clarified the developmental manner of the leaf ofBotrychium ternatum (Thunb.) Sw. The leaf primordium develops as an ordinary appendicular leaf which shows a typical hyponastic growth curvature, though it soon takes a conical shape with a tetrahedral apical cell. The leaf primordium forms a sporophyll primordium first, then forms vegetative primary pinnae acting as the trophophyll primordium. The sporophyll primordium also forms sporogenous primary pinnae.The sporophyll initiation begins with establishment of a new apical cell (sporophyll apical cell) near the original leaf apical cell. Through activity of the sporophyll apical cell most of the sporophyll primordium is formed later. In contrast, the vegetative or sporogenous primary pinna begins to develop as a low mound of surface cells on the trophophyll or sporophyll primordium respectively. The apical cell is later established on the summit of each pinna primordium. In the developmental point of view, the sporophyll primordium is not equivalent to the primary pinnae on the trophophyll primordium. The sporophyll may not represent two fused basal pinnae of a leaf, but represents an organ independent of and equivalent to the trophophyll.

Developmental study onHypolepis punctata (Thunb.) Mett.

The origins of the first and second petiolar buds ofHypolepis punctata were clarified in relation to the early development of the leaf primordium, which arises from a group of superficial cells of the shoot apical meristem. One of these superficial cells produces a two-sided leaf apical cell which subsequently cuts off segments to make a well-defined cell group, called here the leaf apical cell complex, on the distal part of the leaf primordium. Meanwhile, cells surrounding the leaf apical cell complex also divide frequently to form the basal part of the leaf primordium.Two groups of basal cells of the leaf primordium located on the abaxial and the adaxial sides initiate the first and the second petiolar buds, respectively. The initial cells are usually contiguous to the leaf apical cell complex, constructing the abaxial and adaxial flanks of the very young leaf primordium. However, the first petiolar bud sometimes develops from cells located farther from the leaf apical cell complex. These cells are derived from those originally situated in the peripheral region of the shoot apical meristem.

Comparative development of gametophytes ofOsmunda lancea andO. japonica (osmundaceae): Adaptation of rheophilous fern gametophyte

Development of the gametophytes of a rehophytic fernOsmunda lancea and a related dryland speciesO. japonica was examined, and adaptation ofO. lancea gametophytes to rheophytic habitat was discussed. The spores ofO. lancea were larger in size and contained larger oil droplets than those ofO. japonica. The germination rate was similar. In agar culture,O. lancea gametophytes grew more rapidly and reached reprudctive maturity earlier thanO. japonica, although there was no prominent gross-morphological difference in developing and mature gametophytes. Sporophyte formation ofO. lancea took place at roughly two times higher rate than that ofO. japonica. TheO. lancea gametophytes might be adapted to the periodically flooded habitat by shortening its life, which is a different strategy from that of the sporophytic generation.

Surface-viewed shoot apex ofAngiopteris lygodiifolia Ros. (Marattiaceae)

Marattian ferns are thought to be an exception to the rule that a single apical cell is always present in the shoot apex of ferns; the occurrence of plural apical initials has been generally accepted for these ferns. However, a contradicting conclusion was reached in this study which examined the apical organization of the shoot ofAngiopteris lygodiifolia Ros., using fresh materials which had not been fixed. Shoot apices were hand-sectioned transversely into thin sections, including the surface layer of the shoot apex, which were observed by differential interference contrast microscopy without staining.In contrast with the generally accepted view, the shoot apex ofA. lygodiifolia was found to usually possess a single apical cell with three cutting faces. The segments cut off from the apical cell are regularly arranged in a helical sequence. The apical cell seems to actually function as an initial cell of the whole shoot apex. The shoot apices, particularly those of plants cultivated in a greenhouse, sometimes show somewhat irregular organization. In extreme cases, no apical cell is recognizable. However, even in these exceptional cases of such apparently irregular shoot apices, plural apical initials are not found.