Part 10

291. But anthers may become _one-celled_, and that either by confluence or by suppression.

292. By confluence, when the two cells run together into one, as they nearly do in most species of Pentstemon (Fig. 297), more so in Monarda (Fig. 300), and completely in the Mallow (Fig. 298) and all the Mallow family.

293. By suppression in certain cases the anther may be reduced to one cell or halved. In Globe Amaranth (Fig. 299) there is a single cell without vestige of any other. Different species of Sage and of the White Sages of California show various grades of abortion of one of the anther-cells, along with a singular lengthening of the connective (Fig. 302-305).

294. The splitting open of an anther for the discharge of its pollen is termed its _Dehiscence_.

295. As the figures show, this is commonly by a line along the whole length of each cell, either lateral or, when the anthers are extrorse, often along the outer face, and when introrse, along the inner face of each cell. Sometimes the opening is only by a chink, hole, or pore at the top, as in the Azalea, Pyrola (Fig. 307), etc.; sometimes a part of the face separates as a sort of trap-door (or valve), hinged at the top, and opening to allow the escape of the pollen, as in the Sassafras, Spice-bush, and Barberry (Fig. 308).

296. =Pollen.= This is the powdery matter, commonly of a yellow color, which fills the cells of the anther, and is discharged during blossoming, after which the stamens generally fall or wither away. Under the microscope it is found to consist of grains, usually round or oval, and all alike in the same species, but very different in different plants. So that the plant may sometimes be recognized from the pollen alone. Several forms are shown in the accompanying figures.

297. An ordinary pollen-grain has two coats; the outer coat thickish, but weak, and frequently adorned with lines or bands, or studded with points; the inner coat is extremely thin and delicate, but extensible, and its cavity when fresh contains a thickish protoplasmic fluid, often rendered turbid by an immense number of minute particles that float in it. As the pollen matures this fluid usually dries up, but the protoplasm does not lose its vitality. When the grain is wetted it absorbs water, swells up, and is apt to burst, discharging the contents. But when weak syrup is used it absorbs this slowly, and the tough inner coat will sometimes break through the outer and begin a kind of growth, like that which takes place when the pollen is placed upon the stigma.

298. Some pollen-grains are, as it were, lobed (as in Fig. 315, 316), or formed of four grains united (as in the Heath family, Fig. 317): that of Pine (Fig. 318) has a large rounded and empty bladder-like expansion upon each side. This renders such pollen very buoyant, and capable of being transported to a great distance by the wind.

299. In species of Acacia simple grains lightly cohere into globular pellets. In Milkweeds and in most Orchids all the pollen of an anther-cell is compacted or coherent into one mass, called a _Pollen-mass_, or POLLINIUM, plural POLLINIA. (Fig. 319-322.)

Section X. PISTILS IN PARTICULAR.

Sec. 1. ANGIOSPERMOUS OR ORDINARY GYNOECIUM.

300. =Gynoecium= is the technical name for the pistil or pistils of a flower taken collectively, or for whatever stands in place of these. The various modifications of the gynoecium and the terms which relate to them require particular attention.

301. The PISTIL, when only one, occupies the centre of the flower; when there are two pistils, they stand facing each other in the centre of the flower; when several, they commonly form a ring or circle; and when very numerous, they are generally crowded in rows or spirals on the surface of a more or less enlarged or elongated receptacle. Their number gives rise to certain terms, the counterpart of those used for stamens (284), which are survivals of the names of orders in the Linnaean artificial system. The names were coined by prefixing Greek numerals to _-gynia_ used for gynoecium, and changed into adjectives in the form of _-gynous_. That is, a flower is

_Monogynous_, when it has a single pistil, whether that be simple or compound;

_Digynous_, when it has only two pistils; _Trigynous_, when with three; _Tetragynous_, with four; _Pentagynous_, with five; _Hexagynous_, with six; and so on to _Polygynous_, with many pistils.

302. =The Parts of a Complete Pistil=, as already twice explained (16, 236), are the OVARY, the STYLE, and the STIGMA. The ovary is one essential part: it contains the rudiments of seeds, called OVULES. The stigma at the summit is also essential: it receives the pollen, which fertilizes the ovules in order that they may become seeds. But the style, commonly a tapering or slender column borne on the summit of the ovary, and bearing the stigma on its apex or its side, is no more necessary to a pistil than the filament is to the stamen. Accordingly, there is no style in many pistils: in these the stigma is _sessile_, that is, rests directly on the ovary (as in Fig. 326). The stigma is very various in shape and appearance, being sometimes a little knob (as in the Cherry, Fig. 271), sometimes a point or small surface of bare tissue (as in Fig. 327-330), and sometimes a longitudinal crest or line (as in Fig. 324, 341-343), or it may occupy the whole length of the style, as in Fig. 331.

303. The word Pistil (Latin, _Pistillum_) means a pestle. It came into use in the first place for such flowers as those of Crown Imperial, or Lily, in which the pistil in the centre was likened to the pestle, and the perianth around it to the mortar, of the apothecary.

304. A pistil is either _simple_ or _compound_. It is simple when it answers to a single flower-leaf, compound when it answers to two or three, or a fuller circle of such leaves conjoined.

305. =Carpels.= It is convenient to have a name for each flower-leaf of the gynoecium; so it is called a _Carpel_, in Latin _Carpellum_ or _Carpidium_. A simple pistil is a carpel. Each component flower-leaf of a compound pistil is likewise a carpel. When a flower has two or more pistils, these of course are simple pistils, that is, separate carpels or pistil-leaves. There may be only a single simple pistil to the flower, as in a Pea or Cherry blossom (Fig. 271); there may be two such, as in many Saxifrages; or many, as in the Strawberry. More commonly the single pistil in the centre of a blossom is a compound one. Then there is seldom much difficulty in ascertaining the number of carpels or pistil-leaves that compose it.

306. =The Simple Pistil=, viewed morphologically, answers to a leaf-blade with margins incurved and united where they meet, so forming a closed case or pod (the ovary), and bearing ovules at the suture or junction of these margins: a tapering upper portion with margins similarly inrolled, is supposed to form the style; and these same margins, exposed at the tip or for a portion of the length, become the stigma. Compare, under this view, the three accompanying figures.

307. So a simple pistil should have a one-celled ovary, only one line of attachment for the ovules, a single style, and a single stigma. Certain variations from this normal condition which sometimes occur do not invalidate this morphological conception. For instance, the stigma may become two-lobed or two-ridged, because it consists of two leaf-margins, as Fig. 324 shows; it may become 2-locellate by the turning or growing inward of one of the sutures, so as to divide the cavity.

308. There are two or three terms which primarily relate to the parts of a simple pistil or carpel, and are thence carried on to the compound pistil, viz.:--

VENTRAL SUTURE, the line which answers to the united margins of the carpel-leaf, therefore naturally called a suture or seam, and the ventral or inner one, because in the circle of carpel-leaves it looks inward or to the centre of the flower.

DORSAL SUTURE is the line down the back of the carpel, answering to the midrib of the leaf,--not a seam therefore; but at maturity many fruits, such as pea-pods, open by this dorsal as well as by the ventral line.

PLACENTA, a name given to the surface, whatever it be, which bears the ovules and seeds. The name may be needless when the ovules grow directly on the ventral suture, or from its top or bottom; but when there are many ovules there is usually some expansion of an ovule-bearing or seed-bearing surface; as is seen in our Mandrake or Podophyllum, Fig. 326.

309. =A Compound Pistil= is a combination of two, three, or a greater number of pistil-leaves or carpels in a circle, united into one body, at least by their ovaries. The annexed figures should make it clear. A series of Saxifrages might be selected the gynoecium of which would show every gradation between two simple pistils, or separate carpels, and their complete coalescence into one compound and two-celled ovary. Even when the constituent styles and stigmas are completely coalescent into one, the nature of the combination is usually revealed by some external lines or grooves, or (as in Fig. 328-330) by the internal partitions, or the number of the placentae. The simplest case of compound pistil is that

310. =With two or more Cells and Axile Placentae=, namely, with as many cells as there are carpels, that have united to compose the organ. Such a pistil is just what would be formed if the simple pistils (two, three, or five in a circle, as the case may be), like those of a Paeony or Stonecrop (Fig. 224, 225), pressed together in the centre of the flower, were to cohere by their contiguous parts. In such a case the placentae are naturally _axile_, or all brought together in the axis or centre; and the ovary has as many DISSEPIMENTS, or internal _Partitions_, as there are carpels in its composition. For these are the contiguous and coalescent walls or sides of the component carpels. When such pistils ripen into pods, they often separate along these lines into their elementary carpels.

311. =One-celled, with free Central Placenta.= The commoner case is that of Purslane (Fig. 272) and of the Pink and Chickweed families (Fig. 331, 332). This is explained by supposing that the partitions (such as those of Fig. 329) have early vanished or have been suppressed. Indeed, traces of them may often be detected in Pinks. On the other hand, it is equally supposable that in the Primula family the free central is derived from parietal placentation by the carpels bearing ovules only at base, and forming a consolidated common placenta in the axis. Mitella and Dionaea help out this conception.

312. =One-celled, with Parietal Placentae.= In this not uncommon case it is conceived that the two or three or more carpel-leaves of such a compound pistil coalesce by their adjacent edges, just as sepal-leaves do to form a gamosepalous calyx, or petals to form a gamopetalous corolla, and as is shown in the diagram, Fig. 333, and in an actual cross-section, Fig. 334. Here each carpel is an open leaf, or with some introflexion, bearing ovules along its margins; and each placenta consists of the contiguous margins of two pistil-leaves grown together. There is every gradation between this and the three-celled ovary with the placentae in the axis, even in the same genus, sometimes even in different stages in the same pistil (Fig. 335, 336).

Sec. 2. GYMNOSPERMOUS GYNOECIUM.

313. The ordinary pistil has a closed ovary, and accordingly the pollen can act upon the contained ovules only indirectly, through the stigma. This is expressed in a term of Greek derivation, viz.:--

_Angiospermous_, meaning that the seeds are borne in a sac or closed vessel. The counterpart term is

_Gymnospermous_, meaning naked-seeded. This kind of pistil, or gynoecium, the simplest of all, yet the most peculiar, characterizes the Pine family and its relatives.

314. While the ordinary simple pistil is conceived by the botanist to be a leaf rolled together into a closed pod (306), those of the Pine, Larch (Fig. 337), Cedar, and Arbor-Vitae (Fig. 338, 339) are open leaves, in the form of scales, each bearing two or more ovules on the inner face, next the base. At the time of blossoming, these pistil-leaves of the young cone diverge, and the pollen, so abundantly shed from the staminate blossoms, falls directly upon the exposed ovules. Afterward the scales close over each other until the seeds are ripe. Then they separate that the seeds may be shed. As the pollen acts directly on the ovules, such pistil (or organ acting as pistil) has no stigma.

315. In the Yew, and in Torreya and Gingko, the gynoecium is reduced to extremest simplicity, that is, to a naked ovule, without any visible carpel.

316. In Cycas the large naked ovules are borne on the margins or lobes of an obvious open leaf. All GYMNOSPERMOUS plants have other peculiarities, also distinguishing them, as a class, from ANGIOSPERMOUS plants.

Section XI. OVULES.

317. =Ovule= (from the Latin, meaning a little egg) is the technical name of that which in the flower answers to and becomes the seed.

318. Ovules are _naked_ in gymnospermous plants (as just described), in all others they are enclosed in the ovary. They may be produced along the whole length of the cell or cells of the ovary, and then they are apt to be numerous, or only from some part of it, generally the top or the bottom. In this case they are usually few or single (_solitary_, as in Fig. 341-343). They may be _sessile_, i. e. without stalk, or they may be attached by a distinct stalk, the FUNICLE or FUNICULUS (Fig. 340).

319. Considered as to then position and direction in the ovary, they are

_Horizontal_, when they are neither turned upward nor downward, as in Podophyllum (Fig. 326),

_Ascending_, when rising obliquely upwards, usually from the side of the cell, not from its very base, as in the Buttercup (Fig. 341), and the Purslane (Fig. 272),

_Erect_, when rising upright from the very base of the cell, as in the Buckwheat (Fig. 342),

_Pendulous_, when hanging from the side or from near the top, as in the Flax (Fig. 270), and

_Suspended_, when hanging perpendicularly from the very summit of the cell, as in the Anemone (Fig. 343). All these terms equally apply to seeds.

320. In structure an ovule is a pulpy mass of tissue, usually with one or two coats or coverings. The following parts are to be noted, viz.--

KERNEL or NUCLEUS, the body of the ovule. In the Mistletoe and some related plants, there is only this nucleus, the coats being wanting.

TEGUMENTS, or coats, sometimes only one, more commonly two. When two, one has been called PRIMINE, the other SECUNDINE. It will serve all purposes to call them simply outer and inner ovule coats.

ORIFICE, or FORAMEN, an opening through the coats at the organic apex of the ovule. In the seed it is _Micropyle_.

CHALAZA, the place where the coats and the kernel of the ovule blend.

HILUM, the place of junction of the funiculus with the body of the ovule.

321. =The Kinds of Ovules.= The ovules in their growth develop in three or four different ways and thereby are distinguished into

_Orthotropous_ or _Straight_, those which develop without curving or turning, as in Fig. 344. The chalaza is at the insertion or base, the foramen or orifice is at the apex. This is the simplest, but the least common kind of ovule.

_Campylotropous_ or _Incurved_, in which, by the greater growth of one side, the ovule curves into a kidney-shaped outline, so bringing the orifice down close to the base or chalaza; as in Fig. 345.

_Amphitropous_ or _Half Inverted_, Fig. 346. Here the forming ovule, instead of curving perceptibly, keeps its axis nearly straight, and, as it grows, turns round upon its base so far as to become transverse to its funiculus, and adnate to its upper part for some distance. Therefore in this case the attachment of the funiculus or stalk is about the middle, the chalaza is at one end, the orifice at the other.

_Anatropous_ or _Inverted_, as in Fig. 347, the commonest kind, so called because in its growth it has as it were turned over upon its stalk, to which it has continued adnate. The organic base, or chalaza, thus becomes the apparent summit, and the orifice is at the base, by the side of the hilum or place of attachment. The adnate portion of the funiculus, which appears as a ridge or cord extending from the hilum to the chalaza, and which distinguishes this kind of ovule, is called the RHAPHE. The amphitropous ovule (Fig. 346) has a short or incomplete rhaphe.

322. Fig. 348-352 show the stages through which an ovule becomes anatropous in the course of its growth. The annexed two figures are sections of such an ovule at maturity; and Fig. 355 is Fig. 353 enlarged, with the parts lettered.

Section XII. MODIFICATIONS OF THE RECEPTACLE.

323. =The Torus= or Receptacle of the flower (237, Fig. 223) is the portion which belongs to the stem or axis. In all preceding illustrations it is small and short. But it sometimes lengthens, sometimes thickens or variously enlarges, and takes on various forms. Some of these have received special names, very few of which are in common use. A lengthened portion of the receptacle is called