Ovules naked, rarely without carpellary leaves, usually borne on carpophylls, which assume various forms. The single megaspore enclosed in the nucellus is filled with tissue (prothallus) before fertilization, and contains two or more archegonia, consisting usually of a large egg-cell and a small neck, rarely of an egg-cell only and no neck (Gnetum and Welwitschia).
In the genus Cycas the female flower is peculiar among cycads in consisting of a terminal crown of separate leaf-like carpels several inches in length; the apical portion of each carpellary leaf may be broadly triangular in form, and deeply dissected on the margins into narrow woolly appendages like rudimentary pinnae.
In Cycas the stem apex, after producing a cluster of carpellary leaves, continues to elongate and produces more budscales, which are afterwards pushed aside as a fresh crown of fronds is developed.
The various genera are distinguished from one another by the shape and manner of attachment of the pinnae, the form of the carpellary scales, and to some extent by anatomical characters.
- Large cones; the carpellary scales terminate in a peltate distal expansion.
In structure a cycadean sporangium recalls those of certain ferns (Marattiaceae, Osmundaceae and Schizaeaceae), but in the development of the spores there are certain peculiarities not met with among the Vascular Cryptogams. With the exception of Cycas, the female flowers are also in the form of cones, bearing numerous carpellary scales.
Cases have been recorded (by Thiselton-Dyer in Encephalartos and by Wieland in Zamia) in which the short carpellary cone-scales exhibit a foliaceous form.
As the cone grows in size and becomes woody the lower half of the cone-scale, which we may call the carpellary scale, may remain small, and is so far outgrown by the upper half (seminiferous scale) that it is hardly recognizable in the mature cone.
Abies pectinata, &c.) the ripe cone differs from those of Pinus, Picea and Cedrus in the large size of the carpellary scales, which project as conspicuous thin appendages beyond the distal margins of the broader and more woody seminiferous scales; the long carpellary scale is a prominent feature also in the cone of the Douglas pine (Pseudotsuga Douglasii).
The female flowers of the Taxaceae assume another form; in Microcachrys (Tasmania) the reproductive structures are spirally disposed, and form small globular cones made up of red fleshy scales, to each of which is attached a single ovule enclosed by an integument and partially invested by an arillus; in Dacrydium the carpellary leaves are very similar to the foliage leaves - each bears one ovule with two integuments, the outer of which constitutes an arillus.
It is important to draw attention to some structural features exhibited by certain cone-scales, in which there is no external sign indicative of the presence of a carpellary and a seminiferous scale.
The seminiferous scale of Pinus, &c., is also spoken of sometimes as a ligular outgrowth from the carpellary leaf.
Another view is to regard the cone as an inflorescence, each carpellary scale being a bract bearing in its axil a shoot the axis of which has not been developed; the seminiferous scale is believed to represent either a single leaf or a fused pair of leaves belonging to the partially suppressed axillary shoot.
In 1869 van Tieghem laid stress on anatomical evidence as a key to the morphology of the cone-scales; he drew attention to the fact that the collateral vascular bundles of the seminiferous scale are inversely orientated as compared with those of the carpellary scale; in the latter the xylem of each bundle is next the upper surface, while in the seminiferous scale the phloem occupies that position.
In a young cone the seminiferous scale appears as a hump of tissue at the base or in the axil of the carpellary scale, but Celakovsky, a strong supporter of the axillary-bud theory, attaches little or no importance to this kind of evidence, regarding the present manner of development as being merely an example of a short cut adopted in the course of evolution, and replacing the original production of a branch in the axil of each carpellary scale.
Eichler, one of the chief supporters of the simpler view, does not recognize in the inverse orientation of the vascular bundles an argument in support of the axillary-bud theory, but points out that the seminiferous scale, being an outgrowth from the surface of the carpellary scale, would, like outgrowths from an ordinary leaf, naturally have its bundles inversely orientated.
15, D) Sequoia, &c., there are always two sets of bundles; the upper set, having the phloem uppermost, as in the seminiferous scale of Abies or Pinus, are regarded as belonging to the outgrowth from the carpellary scale and specially developed to supply the ovules.
The occurrence of buds in the axils of carpellary scales may, however, simply mean that buds, which are (C and D after Worsdell.) FIG.
(In rare cases the proliferated portion produces male flowers in the leaf -axils.) In Larix the carpellary scale may become leafy, and the seminiferous scale may disappear.
16), in which the lower part bears stamens and the upper portion carpellary and seminiferous scales.
Without expressing any decided opinion as to the morphology of the double cone-scale of the Abietineae, preference may be felt in favour of regarding the cone-scale of the Araucarieae as a simple carpellary leaf bearing a single ovule.
The seeds of some genera depend on animals for dispersal, the carpellary scale (Microcachrys) or the outer integument being brightly coloured and attractive.
In many cases, however, the placentas are formations from the axis (axile), and are not connected with the carpellary leaves.
In marginal placentation the part of the carpel bearing the placenta is the inner or ventral suture, corresponding to the margin of the folded carpellary leaf, while the outer or dorsal suture corresponds to the midrib of the carpellary leaf.
- Diagrammatic section of a five-carpellary ovary, in which the edges of the carpels, bearing the placentas and ovules o, are not folded inwards.
- Diagrammatic section of a five-carpellary ovary, in which the septa (s) proceed inwards for a certain length, bearing the placentas and ovules (o).
97), Pyrola, &c. The carpellary leaves.
Carpellary leaves, and that in the progress of development these leaves separated from them, leaving the placentas and ovules free in the centre; or by supposing that the placentas are not marginal but axile formations, produced by an elongation of the axis, and the carpels verticillate leaves, united together around the axis.
The carpellary leaves, but are also formed over the whole surface of the leaf, as in Butomus.
In Cycas the altered leaf, upon the margin of which the ovule is produced, and the peltate scales, from which they are pendulous in Zamia, are regarded by all botanists as carpellary leaves.
The carpellary leaves are sometimes united in such a way as to leave an opening at the apex of the pistil, so that the ovules are exposed, as in mignonette.