Part 42

The reduction in the ANGIOSPERMS is carried to the extreme limit. In the embryo-sac (the macrospore) the nucleus by continued division produces a prothallium consisting of primordial cells (Fig. 254). In the upper end of the embryo-sac (which is nearest the micropyle) are three cells, two of which are termed the “co-operating cells” (synergidæ) and the third is the oosphere. Three others are placed at the opposite end of the embryo-sac and are therefore termed the “antipodal cells.” Finally, a large cell is also formed, which occupies the space between the two groups and whose cell-nucleus, the central definitive nucleus, lies in the centre of the embryo-sac. These primordial cells are the slight remnant of the prothallium. The entire structure of the archegonium, with its neck and canal-cells, has disappeared, and nothing is left but the indispensable oosphere. When the oosphere has been fertilised, and has commenced the cellular divisions which lead to the formation of the embryo (Fig. 255), the synergidæ and antipodal cells are absorbed, and a cell-formation begins by a new process which emanates from the definitive nucleus and by which a parenchymatous cell-tissue, the nutritive-tissue, arises which may perhaps be considered as homologous with the endosperm of the Gymnosperms. The difference is that the nutritive-tissue of the Angiosperms is formed in two parts with an intervening interruption; the primary nutritive-tissue is first formed, and after fertilisation is absorbed, with the exception of one cell, which continues the development and gives rise to the nutritive-tissue proper, which is formed in the first instance of primordial cells, and later on of a cellular tissue; this nutritive-tissue formed in the embryo-sac is termed “endosperm”; in a few instances a tissue which is derived from the nucellus functions as nutritive-tissue, and is termed “perisperm.” In many plants the seeds, when ripe, contain a very rich nutritive-tissue, in addition to the embryo, for the purpose of its nourishment during germination. These are termed albuminous (endospermous) seeds, in distinction to the ex-albuminous, or those in which the nutritive-tissue is stored in the embryo itself, before it is completely developed, and used for its sustenance.

In addition to the changes which fertilisation produces in the ovule itself, it also gives the impetus to a series of changes in the entire shoot which bears the ovule. The perianth, stamens, and style, generally wither, because the part they play is at an end; the wall of the ovary grows and becomes the wall of the fruit (pericarp). The entire gynœcium of a flower, transformed as a consequence of fertilisation, is termed a fruit. It consists of two parts, the pericarp and the seeds, and according to the nature of the pericarp, the fruit is termed a capsule, nut, berry, or drupe.

The chief characteristic of the Phanerogams does not lie in the formation of the flower (although they may quite properly be termed “Flowering-plants”), because Equisetums and Lycopods have reproductive shoots as highly differentiated as those of certain Gymnosperms and other Phanerogams. As regards the SEXUAL GENERATION the characteristics are found:--(1) in its great reduction; (2) in the transmission of the microspore (pollen-grain) to the macrosporangium, and its germination, with the formation of a pollen-tube (antheridium), the protoplasm of which is not differentiated into spermatozoids; (3) in the fact that the macrospore (embryo-sac) never leaves its sporangium (nucellus); and further in the Angiosperms, (4) in the peculiar development of the nutritive-tissue in two parts; and (5) in the great reduction of the archegonium.

As regards the ASEXUAL GENERATION the characteristic feature is that this generation is formed whilst the sporangium is still attached to the mother-plant, and for a long time is nourished by it; and that after the sporangium has become detached from the mother-plant, it spends a longer or shorter resting period as the embryo in the seed (enveloped by the testa), and does not make its appearance until the “germination” of the seed. In addition the shoot which bears sporangia undergoes greater modification than in the case of the Flowerless-plants.

The Phanerogams are separated into two Divisions as follows:--

Division 4. =Gymnospermæ.= The ovules, as well as the seeds, are borne naked on the surface of open carpels, or on the apex of a stem (ovary wanting). The pollen-grains are conveyed by the wind to the ovules, and caught by drops of mucilage, secreted by the micropyle. A “stigma” is wanting. The entire female prothallium (the endosperm), which serves for the nourishment of the embryo, is formed before fertilisation. The archegonia are embedded in the upper part of the prothallium. The pollen-grains are “multicellular,” i.e. there is always in their interior a distinct prothallium, formed by 1–3 cells, and a larger cell which gives rise to the pollen-tube.

Division 5. =Angiospermæ.= The carpels surround the ovules and form an entirely closed chamber (ovary), in which the ovules mature and ripen into seeds. The surface of a portion of the apex of the carpel is transformed into the “stigma,” which, by a sticky fluid and also by hair-structures, is capable of retaining the pollen-grains conveyed to it by the wind, or more frequently by insects. The pollen-tube grows from the stigma, through the “conducting cellular tissue” (style), to the ovules. The pollen-grains contain two cells, a vegetative and a free generative cell. The latter passes into the pollen-tube and there divides into two, one of which is the sperm-nucleus. The female prothallium, which is intended to serve as nutritive-tissue, is formed after fertilisation. Archegonia are wanting.

DIVISION IV.

GYMNOSPERMÆ.

The following characters should be added to those already given on page 2:--

The Gymnosperms comprise only trees or shrubs. The flowers are always unisexual and destitute of perianth (except Gnetaceæ); the female plant of Cycas is the only one which has no flower. The MALE FLOWERS are constructed on the same type as the cones of the Horsetails and Club-Mosses, and are most frequently long shoots (Figs. 243, 258, 260 A, 267 J) bearing a number of spiral or verticillate stamens. The FEMALE FLOWERS are of a more varied structure (see the orders). The OVULE is orthotropous (except Podocarpus which is anatropous) and projects from the carpel uprightly, inverted, or horizontally; it has usually only one integument (compare however Taxaceæ) which proceeds from the upper part of the nucellus, so that the embryo-sac in part is placed below the integuments (Figs. 251, 264). The drop of mucilage which catches the pollen-grains dries up and draws the pollen-grain through the micropyle to a space just above the nucellus--the pollen-chamber--in which the germination of the pollen-grain commences.

In each seed, only one of the many embryos which are formed proceeds to its full development. The seed is always endospermous, and the embryo has one, two, or a whorl of several cotyledons. A vigorous primary root is developed on germination. THE VASCULAR BUNDLES in the stem are arranged in a ring, and increase in thickness takes place by a closed cambium-ring which forms bast (phlœem) externally, and wood (xylem) internally with distinct annual rings, as in the Dicotyledons. Only certain of the Cycadeæ deviate from this arrangement. The secondary wood is very uniform, as it is almost exclusively formed of tracheides with bordered pits, but true vessels are wanting; this also indicates a relationship with the Pteridophyta (see page 202).

The Gymnosperms are biologically lower than the Angiosperms; they are wind-fertilised, and without extra floral-nectaries.

This Division embraces three classes: CYCADEÆ, CONIFERÆ, And GNETEÆ. It is no doubt monophyletic, and has taken its origin from heterosporous Ferns, now extinct, most nearly related to the Ophioglossaceæ and Marattiaceæ. The Cycadeæ appear to be the oldest class. The Coniferæ are related to these through Ginkgo. The Gnetaceæ are more isolated. The Division is not continued into the higher Flowering-plants; it has evidently attained its highest development, and is now in a retrograde condition. The similarity which has often been pointed out between certain Coniferæ and Lycopodinæ is only in analogous resemblances, and does not entitle one to suppose that there is a nearer relationship, or that the former take their origin from the latter.

Class 1. =Cycadeæ.=

The stem is very rarely ramified. The leaves are large, pinnate, and arranged spirally. The flowers are diœcious, without perianth.

There is only one order, the =Cycadaceæ=.--In habit they resemble the Ferns, especially the Tree-Ferns (compare Figs. 207 and 256). The stem is tubercular (Fig. 258), or cylindrical (Fig. 256), but not very tall (as much as about 12 metres), and very rarely ramified. [In Ceylon, unbranched specimens of Cycas are rarely met with in the wild state. The stems of C. circinalis occasionally branch in greenhouses.]

The LEAVES are arranged spirally, and so closely together that no free stem-surface is left between them, and have only a slight sheath (which is not amplexicaul, as in the Palms). They are compound (most frequently pinnate; in Bowenia, bipinnate); in some genera the leaves are rolled up in various ways, resembling the vernation in Ferns (Fig. 257); they are leathery and perennial. In some, stipules are present, as in the Marattiaceæ. Groups of scale-leaves alternate in the majority with groups of foliage-leaves.

The FLOWERS are without perianth. The MALE FLOWER is sometimes an enormous collection of stamens (Fig. 258), which are flat in some (Cycas, Fig. 241), shield-like in others (Zamia, Ceratozamia) like the sporophylls in Horsetail (Fig. 259); but in all, the pollen-sacs are situated in large and varying numbers on the back of the stamens, and arranged in groups of 2–5, like the sporangia in the sori of the Ferns (Fig. 241 b, c). FEMALE FLOWERS are wanting in Cycas, because the carpels do not terminate the apical growth of the stem. After a group of foliage-and of scale-leaves, a group of carpels is developed, which are pinnate and resemble the foliage-leaves, bearing on their edges a number of ovules (most frequently 5–6) (Figs. 245, 256); the same stem produces successively scale-leaves, foliage-leaves, and carpels. The differentiation is not much more advanced than in certain Ferns (Struthiopteris, Blechnum), where barren and fertile leaves of different form regularly alternate. The other genera have female flowers; the carpels are shield-like in Zamia and Ceratozamia (Fig. 246), and collected into cone-like flowers, which terminate the growth of the stem (Fig. 259). The number of ovules in these instances is two to each carpel.

The SEEDS are large (most frequently 2–6 centimetres long) and plum-like; the external layer of the testa is fleshy, while the internal one is hard and horny. There are two systems of vascular bundles in the testa, one outside, the other inside the stone. The embryo is straight, attached to the end of the suspensor, which is often long, filamentous, and rolled up; it has one or two cotyledons.

The embryo in Ceratozamia and others is very slightly developed, at the time when the ripe seed is detached from the carpel; and it is not until after sowing that its further development and germination proceed. This calls to mind the Cryptogams, especially Selaginella, whose macrospores are thrown off filled with endosperm; but the oosphere is not fertilised till after the separation of the macrospore from the parent-plant, while in the Cycadeæ fertilisation is effected before the separation. In Cycas the testa may rupture, and the endosperm grow and become green in the light, even though no embryo has been formed. This also is an indication of its prothalloid nature.

Gum-passages are present in all organs. Collateral vascular bundles, with spiral and scalariform tracheides, are found; and normal thickening takes place by means of a cambium. An exceptional mode of growth is found in Cycas and Encephalartos, the cambium ceases to divide after a time and is replaced by a new cambium which arises in the cortical parenchyma just outside the bast, and which forms a new ring of xylem and phlœem. This may be repeated so that a number of concentric rings are produced. In Ceratozamia, structures resembling corals extend from the roots in a vertical direction and appear on the surface of the soil; these are peculiar roots, in which a symbiotic Alga (Anabæna) is found.

The Cycadeæ were formerly (from the Coal period to the Later Cretaceous) far more numerous than at the present day. They appear to have been most numerous in the Trias and Jurassic. The remnant (75 species) which have persisted to the present time are found in all tropical countries. Cycas (Trop. and Sub-trop., Eastern Hemisphere); Dioon (Mexico); Macrozamia (Australia); Encephalartos (Trop. and S. Africa); Stangeria (Fig. 258, Sub-trop. South and East Africa); Bowenia (Trop. Australia); Ceratozamia (Mexico, New Granada, Western Brazil); Microcycas (Cuba); Zamia (Trop. and Sub-trop. N. America.)

USES. Sago is made from the starch-containing pith of Cycas revoluta and circinalis. The leaves are often used at funerals and church festivals, under the name of “palm-branches.”

Class 2. =Coniferæ= (=Pine-trees=).

The stem branches freely. The leaves are entire, relatively small, linear or reduced to scales. The flowers are without perianth. The ovules naked. It is seldom that the female flower is reduced to only one carpel.

Whilst the Cycadeæ principally resemble the Ferns, the Conifers partly resemble the Lycopods, and partly the Equisetums--the former especially in the needle- or scale-like, leathery, simple, and often perennial leaves (“evergreen plants”), which never possess stipules (Figs. 263, 270, 272). Ginkgo deviates from this, being no doubt the oldest, and the Conifer which stands nearest to the Cycadeæ (Fig. 260). The resemblance to the Equisetums is especially owing to the fact that the stem ramifies abundantly, and often very regularly, forming a pyramid with verticillate branches. In addition to the foliage-leaves, scale-leaves (bud-scales) are present in the majority of species.

The FLOWERS are monœcious or more rarely diœcious. Perianth is wanting. The stamens of the catkin-like male flowers (Fig. 267, J) are of different forms, but as a rule more or less shield-like. As in the Cycadeæ, the pollen-sacs are in all cases situated on the underside. There are, as a rule, two pollen-sacs (the Abietaceæ, Fig. 267), or 3–5, (the Cupressaceæ and Taxaceæ, Fig. 243); a few have more, e.g. Araucaria (Fig. 242); they dehisce by clefts.