(P. McC.)
GRASSE, FRANÇOIS JOSEPH PAUL,Marquis de Grassetilly, Comte de(1722-1788), French sailor, was born at Bar, in the present department of the Alpes Maritimes. In 1734 he took service on the galleys of the order of Malta, and in 1740 entered the service of France, being promoted to chief of squadron in 1779. He took part in the naval operations of the American War of Independence, and distinguished himself in the battles of Dominica and Saint Lucia (1780), and of Tobago (1781). He was less fortunate at St Kitts, where he was defeated by Admiral Hood. Shortly afterwards, in April 1782, he was defeated and taken prisoner by Admiral Rodney. Some months later he returned to France, published aMémoire justificatif, and was acquitted by a court-martial (1784). He died at Paris in January 1788.
His son Alexandre de Grasse, published aNotice bibliographique sur l’amiral comte de Grasse d’après les documents inéditsin 1840. See G. Lacour-Gayet,La Marine militaire de la France sous le règne de Louis XV(Paris, 1902).
His son Alexandre de Grasse, published aNotice bibliographique sur l’amiral comte de Grasse d’après les documents inéditsin 1840. See G. Lacour-Gayet,La Marine militaire de la France sous le règne de Louis XV(Paris, 1902).
GRASSE,a town in the French department of the Alpes Maritimes (till 1860 in that of the Var), 12½ m. by rail N. of Cannes. Pop. (1906) town, 13,958; commune, 20,305. It is built in a picturesque situation, in the form of an amphitheatre and at a height o£ 1066 ft. above the sea, on the southern slope of a hill, facing the Mediterranean. In the older (eastern) part of the town the streets are narrow, steep and winding, but the new portion (western) is laid out in accordance with modern French ideas. It possesses a remarkably mild and salubrious climate, and is well supplied with water. That used for the purpose of the factories comes from the fine spring of Foux. But the drinking water used in the higher portions of the town flows, by means of a conduit, from the Foulon stream, one of the sources of the Loup. Grasse was from 1244 (when the see was transferred hither from Antibes) to 1790 an episcopal see, but was then included in the diocese of Fréjus till 1860, when politically as well as ecclesiastically, the region was annexed to the newly-formed department of the Alpes Maritimes. It still possesses a 12th-century cathedral, now a simple parish church; while an ancient tower, of uncertain date, rises close by near the town hall, which was formerly the bishop’s palace (13th century). There is a good town library, containing the muniments of the abbey of Lérins, on the island of St Honorat opposite Cannes. In the chapel of the old hospital are three pictures by Rubens. The painter J. H. Fragonard (1732-1806) was a native of Grasse, and some of his best works were formerly to be seen here (now in America). Grasse is particularly celebrated for its perfumery. Oranges and roses are cultivated abundantly in the neighbourhood. It is stated that the preparation of attar of roses (which costs nearly £100 per 2 ℔) requires alone nearly 7,000,000 roses a year. The finest quality of olive oil is also manufactured at Grasse.
(W. A. B. C.)
GRASSES,1a group of plants possessing certain characters in common and constituting a family (Gramineae) of the class Monocotyledons. It is one of the largest and most widespread and, from an economic point of view, the most important family of flowering plants. No plant is correctly termed a grass which is not a member of this family, but the word is in common language also used, generally in combination, for many plants of widely different affinities which possess some resemblance (often slight) in foliage to true grasses;e.g.knot-grass (Polygonum aviculare), cotton-grass (Eriophorum), rib-grass (Plantago), scorpion-grass (Myosotis), blue-eyed grass (Sisyrinchium), sea-grass (Zostera). The grass-tree of Australia (Xanthorrhoea) is a remarkable plant, allied to the rushes in the form of its flower, but with a tall, unbranched, soft-woody, palm-like trunk bearing a crown of long, narrow, grass-like leaves and stalked heads of small, densely-crowded flowers. In agriculture the word has an extended signification to include the various fodder-plants, chiefly leguminous, often called “artificial grasses.” Indeed, formerlygrass(also speltgwrs,gres,gyrsin the old herbals) meant any green herbaceous plant of small size.
Yet the first attempts at a classification of plants recognized and separated a group ofGramina, and this, though bounded by nothing more definite than habit and general appearance, contained the Gramineae of modern botanists. The older group, however, even with such systematists as Ray (1703), Scheuchzer (1719), and Micheli (1729), embraced in addition the Cyperaceae(Sedge family), Juncaceae (Rush family), and some other monocotyledons with inconspicuous flowers. Singularly enough, the sexual system of Linnaeus (1735) served to mark off more distinctly the true grasses from these allies, since very nearly all of the former then known fell under his Triandria Digynia, whilst the latter found themselves under his other classes and orders.
I. Structure.—The general type of true grasses is familiar in the cultivated cereals of temperate climates—wheat, barley, rye, oats, and in the smaller plants which make up pastures and meadows and form a principal factor of the turf of natural downs. Less familiar are the grains of warmer climates—rice, maize, millet and sorgho, or the sugar-cane. Still farther removed are the bamboos of the tropics, the columnar stems of which reach to the height of forest trees. All are, however, formed on a common plan.
Root.—Most cereals and many other grasses are annual, and possess a tuft of very numerous slender root-fibres, much branched and of great length. The majority of the members of the family are of longer duration, and have the roots also fibrous, but fewer, thicker and less branched. In such cases they are very generally given off from just above each node (often in a circle) of the lower part of the stem or rhizome, perforating the leaf-sheaths. In some bamboos they are very numerous from the lower nodes of the erect culms, and pass downwards to the soil, whilst those from the upper nodes shrivel up and form circles of spiny fibres.
Stem.—The underground stem or rootstock (rhizome) of perennial grasses is usually well developed, and often forms very long creeping or subterranean rhizomes, with elongated internodes and sheathing scales; the widely-creeping, slender rhizomes in Marram-grass (Psamma),Agropyrum junceum,Elymus arenarius, and other sand-loving plants render them useful as sand-binders. It is also frequently short, with the nodes crowded. The turf-formation, which is characteristic of open situations in cool temperate climates, results from an extensive production of short stolons, the branches and the fibrous roots developed from their nodes forming the dense “sod.” The very large rhizome of the bamboos (fig. 1) is also a striking example of “definite” growth; it is much branched, the short, thick, curved branches being given off below the apex of the older ones and at right angles to them, the whole forming a series of connected arched axes, truncate at their ends, which were formerly continued into leafy culms. The rhizome is always solid, and has the usual internal structure of the monocotyledonous stem. In the cases of branching just cited the branches break directly through the sheath of the leaf in connexion with which they arise. In other cases the branches grow upwards through the sheaths which they ultimately split from above, and emerging as aerial shoots give a tufted habit to the plant. Good examples are the oat, cock’s-foot (Dactylis) and other British grasses. This mode of growth is the cause of the “tillering” of cereals, or the production of a large number of erect growing branches from the lower nodes of the young stem. Isolated tufts or tussocks are also characteristic of steppe—and savanna—vegetation and open places generally in the warmer parts of the earth.
The aerial leaf-bearing branches (culms) are a characteristic feature of grasses. They are generally numerous, erect, cylindrical (rarely flattened) and conspicuously jointed with evident nodes. The nodes are solid, a strong plate of tissue passing across the stem, but the internodes are commonly hollow, although examples of completely solid stems are not uncommon (e.g.maize, many Andropogons, sugar-cane). The swollen nodes are a characteristic feature. In wheat, barley and most of the British native grasses they are a development, not of the culm, but of the base of the leaf-sheath. The function of the nodes is to raise again culms which have become bent down; they are composed of highly turgescent tissue, the cells of which elongate on the side next the earth when the culm is placed in a horizontal or oblique position, and thus raise the culm again to an erect position. The internodes continue to grow in length, especially the upper ones, for some time; the increase takes place in a zone at the extreme base, just above the node. The exterior of the culms is more or less concealed by the leaf-sheaths; it is usually smooth and often highly polished, the epidermal cells containing an amount of silica sufficient to leave after burning a distinct skeleton of their structure. Tabasheer is a white substance mainly composed of silica, found in the joints of several bamboos. A few of the lower internodes may become enlarged and sub-globular, forming nutriment-stores, and grasses so characterized are termed “bulbous” (Arrhenatherum,Poa bulbosa, &c.). In internal structure grass-culms, save in being hollow, conform to that usual in monocotyledons; the vascular bundles run parallel in the internodes, but a horizontal interlacement occurs at the nodes. In grasses of temperate climates branching is rare at the upper nodes of the culm, but it is characteristic of the bamboos and many tropical grasses. The branches are strictly distichous. In many bamboos they are long and spreading or drooping and copiously ramified, in others they are reduced to hooked spines. One genus (Dinochloa, a native of the Malay archipelago) is scandent, and climbs over trees 100 ft. or more in height,Olyra latifolia, a widely-spread tropical species, is also a climber on a humbler scale.
Grass-culms grow with great rapidity, as is most strikingly seen in bamboos, where a height of over 100 ft. is attained in from two to three months, and many species grow two, three or even more feet in twenty-four hours. Silicic hardening does not begin till the full height is nearly attained. The largest bamboo recorded is 170 ft., and the diameter is usually reckoned at about 4 in. to each 50 ft.
Leaves.—These present special characters usually sufficient for ordinal determination. They are solitary at each node and arranged in two rows, the lower often crowded, forming a basal tuft. They consist of two distinct portions, the sheath and the blade. The sheath is often of great length, and generally completely surrounds the culm, forming a firm protection for the internode, the younger basal portion of which, including the zone of growth, remains tender for some time. As a rule it is split down its whole length, thus differing from that of Cyperaceae which is almost invariably (Eriosporais an exception) a complete tube; in some grasses, however (species ofPoa,Bromusand others), the edges are united. The sheaths are much dilated inAlopecurus vaginatusand in a species ofPotamochloa, in the latter, an East Indian aquatic grass, serving as floats. At the summit of the sheath, above the origin of the blade, is theligule, a usually membranous process of small size (occasionally reaching 1 in. in length) erect and pressed around the culm. It is rarely quite absent, but may be represented by a tuft of hairs (very conspicuous inPariana). It serves to prevent rain-water, which has run down the blade, from entering the sheath.Melica uniflorahas in addition to the ligule, a green erect tongue-like process, from the line of junction of the edges of the sheath.
The blade is frequently wanting or small and imperfect in the basal leaves, but in the rest is long and set on to the sheath at an angle. The usual form is familiar—sessile, more or less ribbon-shaped, tapering to a point, and entire at the edge. The chief modifications are the articulation of the deciduousblade on to the sheath, which occurs in all the Bambuseae (exceptPlanotia) and inSpartina stricta; and the interposition of a petiole between the sheath and the blade, as in bamboos,Leptaspis,Pharus,Pariana,Lophatherumand others. In the latter case the leaf usually becomes oval, ovate or even cordate or sagittate, but these forms are found in sessile leaves also (Olyra,Panicum). The venation is strictly parallel, the midrib usually strong, and the other ribs more slender. InAnomochloathere are several nearly equal ribs and in some broad-leaved grasses (Bambuseae,Pharus,Leptaspis) the venation becomes tesselated by transverse connecting veins. The tissue is often raised above the veins, forming longitudinal ridges, generally on the upper face; the stomata are in lines in the intervening furrows. The thick prominent veins inAgropyrumoccupy the whole upper surface of the leaf. Epidermal appendages are rare, the most frequent being marginal, saw-like, cartilaginous teeth, usually minute, but occasionally (Danthonia scabra,Panicum serratum) so large as to give the margin a serrate appearance. The leaves are occasionally woolly, as inAlopecurus lanatusand one or twoPanicums. The blade is often twisted, frequently so much so that the upper and under faces become reversed. In dry-country grasses the blades are often folded on the midrib, or rolled up. The rolling is effected by bands of large wedge-shaped cells—motor-cells—between the nerves, the loss of turgescence by which, as the air dries, causes the blade to curl towards the face on which they occur. The rolling up acts as a protection from too great loss of water, the exposed surface being specially protected to this end by a strong cuticle, the majority or all of the stomata occurring on the protected surface. The stiffness of the blade, which becomes very marked in dry-country grasses, is due to the development of girders of thick-walled mechanical tissue which follow the course of all or the principal veins (fig. 2).
Inflorescence.—This possesses an exceptional importance in grasses, since, their floral envelopes being much reduced and the sexual organs of very great uniformity, the characters employed for classification are mainly derived from the arrangement of the flowers and their investing bracts. Various interpretations have been given to these glumaceous organs and different terms employed for them by various writers. It may, however, be considered as settled that the whole of the bodies known as glumes and paleae, and distichously arranged externally to the flower, form no part of the floral envelopes, but are of the nature of bracts. These are arranged so as to formspikelets(locustae), and each spikelet may contain one, as inAgrostis(fig. 3) two, as inAira(fig. 4) three, or a great number of flowers, as inBriza(fig. 5)Triticum(fig. 6); in some species ofEragrostisthere are nearly 60. The flowers are, as a rule, placed laterally on the axis (rachilla) of the spikelet, but in one-flowered spikelets they appear to be terminal, and are probably really so inAnthoxanthum(fig. 7) and in two anomalous genera,AnomochloaandStreptochaeta.
In immediate relation with the flower itself, and often entirely concealing it, is thepaleaorpale(“upper pale” of most systematic agrostologists). This organ (fig. 13, 1) is peculiar to grasses among Glumiflorae (the series to which belong the two families Gramineae and Cyperaceae), and is almost always present, certainOryzeaeandPhalarideaebeing the only exceptions. It is of thin membranous consistence, usually obtuse, often bifid, and possesses no central rib or nerve, but has two lateral ones, one on either side; the margins are frequently folded in at the ribs, which thus become placed at the sharp angles. This structure was formerly regarded as pointing to the fusion of two organs, and the pale was considered by Robert Brown to represent two portions soldered together of a trimerous perianth-whorl, the third portion being the “lower pale.” The pale is now generally considered to represent the single bracteole, characteristic of Monocotyledons, the binerved structure being the result of the pressure of the axis of the spikelet during the development of the pale, as inIrisand others.
The flower with its pale is sessile, and is placed in the axis of another bract in such a way that the pale is exactly opposed to it, though at a slightly higher level. It is this second bract or flowering glume which has been generally called by systematists the “lower pale,” and with the “upper pale” was formerly considered to form an outer floral envelope (“calyx,” Jussieu; “perianthium,” Brown). The two bracts are, however, on different axes, one secondary to the other, and cannot therefore be parts of one whorl of organs. They are usually quite unlike one another, but in some genera (e.g.mostFestuceae) are very similar in shape and appearance.
The flowering glume has generally a more or less boat-shaped form, is of firm consistence, and possesses a well-marked central midrib and frequently several lateral ones. The midrib in a large proportion of genera extends into an appendage termed theawn(fig. 4), and the lateral veins more rarely extend beyond the glume as sharp points (e.g.Pappophorum). The form of the flowering glume is very various, this organ being plastic and extensively modified in different genera. It frequently extends downwards a little on the rachilla, forming with the latter a swollen callus, which is separated from the free portion by a furrow. InLeptaspisit is formed into a closed cavity by the union of its edges, and encloses the flower, the styles projecting through the pervious summit. Valuable characters for distinguishing genera are obtained from the awn. This presents itself variously developed from a mere subulate point to an organ several inches in length, and when complete (as inAndropogoneae,AveneaeandStipeae) consists of two well-marked portions, a lower twisted part and a terminal straight portion,usually set in at an angle with the former, sometimes trifid and occasionally beautifully feathery (fig. 8). The lower part is most often suppressed, and in the large group of thePaniceaeawns of any sort are very rarely seen. The awn may be either terminal or may come off from the back of the flowering glume, and Duval Jouve’s observations have shown that it represents the blade of the leaf of which the portion of the flowering glume below its origin is the sheath; the twisted part (so often suppressed) corresponds with the petiole, and the portion of the glume extending beyond the origin of the awn (very long in some species,e.g.ofDanthonia) with the ligule of the developed foliage-leaf. When terminal the awn has three fibro-vascular bundles, when dorsal only one; it is covered with stomate-bearing epidermis.
The flower with its palea is thus sessile in the axil of a floriferous glume, and in a few grasses (Leersia(fig. 9),Coleanthus,Nardus) the spikelet consists of nothing more, but usually (even in uniflorous spikelets) other glumes are present. Of these the two placed distichously opposite each other at the base of the spikelet never bear any flower in their axils, and are called theemptyorbarren glumes(figs. 3, 8). They are the “glumes” of most writers, and together form what was called the “gluma” by R. Brown. They rarely differ much from one another, but one may be smaller or quite absent (Panicum,Setaria(fig. 10),Paspalum,Lolium), or both be altogether suppressed, as above noticed. They are commonly firm and strong, often enclose the spikelet, and are rarely provided with long points or imperfect awns. Generally speaking they do not share in the special modifications of the flowering glumes, and rarely themselves undergo modification, chiefly in hardening of portions (Sclerachne,Manisuris,Anthephora,Peltophorum), so as to afford greater protection to the flowers or fruit. But it is usual to find, besides the basal glumes, a few other empty ones, and these are in two- or more-flowered spikelets (seeTriticum, fig. 6) at the top of the rhachilla (numerous inLophatherum), or in uniflorous ones (fig. 10) below and interposed between the floral glume and the basal pair.
The axis of the spikelet is frequently jointed and breaks up into articulations above each flower. Tufts or borders of hairs are frequently present (Calamagrostis,Phragmites,Andropogon), and are often so long as to surround and conceal the flowers (fig. 11). The axis is often continued beyond the last flower or glume as a bristle or stalk.
a,b, Barren glumes.
c,c, Fertile glumes, each enclosing one flower with its paled.
Note the zigzag axis (rhachilla) bearing long silky hairs.
Involucresor organs outside the spikelets also occur, and are formed in various ways. Thus inSetaria(fig. 10),Pennisetum, &c., the one or more circles of simple or feathery hairs represent abortive branches of the inflorescence; inCenchrus(fig. 12) these become consolidated, and the inner ones flattened so as to form a very hard globular spiny case to the spikelets. The cup-shaped involucre ofCornucopiais a dilatation of the axis into a hollow receptacle with a raised border. InCynosurus(Dog’s tail) the pectinate involucre which conceals the spikelet is a barren or abortive spikelet. Bracts of a more general character subtending branches of the inflorescence are singularly rare in Gramineae, in marked contrast with Cyperaceae, where they are so conspicuous. They however occur in a whole section ofAndropogon, inAnomochloa, and at the base of the spike inSesleria. The remarkable ovoid involucre ofCoix, which becomes of stony hardness, white and polished (then known as “Job’s tears,”q.v.), is also a modified bract or leaf-sheath. It is closed except at the apex, and contains the female spikelet, the stalks of the male inflorescence and the long styles emerging through the small apical orifice.
Any number of spikelets may compose the inflorescence, and their arrangement is very various. In the spicate forms, with sessile spikelets on the main axis, the latter is often dilated and flattened (Paspalum), or is more or less thickened and hollowed out (Stenotaphrum,Rottboellia,Tripsacum), when the spikelets are sunk and buried within the cavities. Every variety of racemose and paniculate inflorescence obtains, and the number of spikelets composing those of the large kinds is often immense. Rarely the inflorescence consists of very few flowers; thusLygeum Spartum, the most anomalous of European grasses, has but two or three large uniflorous spikelets, which are fused together at the base, and have no basal glumes, but are enveloped in a large, hooded, spathe-like bract.
Flower.—This is characterized by remarkable uniformity. The perianth is represented by very rudimentary, small, fleshy scales arising below the ovary, calledlodicules; they are elongated or truncate, sometimes fringed with hairs, and are in contact with the ovary. Their usual number is two, and they are placed collaterally at the anterior side of the flower (fig. 13,) that is, within the flowering glume. They are generally considered to represent the inner whorl of the ordinary monocotyledonous(liliaceous) perianth, the outer whorl of these being suppressed as well as the posterior member of the inner whorl. This latter is present almost constantly inStipeaeandBambuseae, which have three lodicules, and in the latter group they are occasionally more numerous. InAnomochloathey are represented by hairs. InStreptochaetathere are six lodicules, alternately arranged in two whorls. Sometimes, as inAnthoxanthum, they are absent. InMelicathere is one large anterior lodicule resulting presumably from the union of the two which are present in allied genera. Professor E. Hackel, however, regards this as an undivided second pale, which in the majority of the grasses is split in halves, and the posterior lodicule, when present, as a third pale. On this view the grass-flower has no perianth. The function of the lodicules is the separation of the pale and glume to allow the protrusion of stamens and stigmas; they effect this by swelling and thus exerting pressure on the base of these two structures. Where, as inAnthoxanthum, there are no lodicules, pale and glume do not become laterally separated, and the stamens and stigmas protrude only at the apex of the floret (fig. 7). Grass-flowers are usually hermaphrodite, but there are very many exceptions. Thus it is common to find one or more imperfect (usually male) flowers in the same spikelet with bisexual ones, and their relative position is important in classification.HolcusandArrhenatherumare examples in English grasses; and as a rule in species of temperate regions separation of the sexes is not carried further. In warmer countries monoecious and dioecious grasses are more frequent. In such cases the male and female spikelets and inflorescence may be very dissimilar, as in maize, Job’s tears,Euchlaena,Spinifex, &c.; and in some dioecious species this dissimilarity has led to the two sexes being referred to different genera (e.g.Anthephora axilliflorais the female ofBuchloe dactyloides, andNeurachne paradoxaof a species ofSpinifex). In other grasses, however, with the sexes in different plants (e.g.Brizopyrum,Distichlis,Eragrostis capitala,Gynerium), no such dimorphism obtains.Amphicarpumis remarkable in having cleistogamic flowers borne on long radical subterranean peduncles which are fertile, whilst the conspicuous upper paniculate ones, though apparently perfect, never produce fruit. Something similar occurs inLeersia oryzoides, where the fertile spikelets are concealed within the leaf-sheaths.
Androecium.—In the vast majority there are three stamens alternating with the lodicules, and therefore one anterior,i.e.opposite the flowering glume, the other two being posterior and in contact with the palea (fig. 13, 1 and 2). They are hypogynous, and have long and very delicate filaments, and large, linear or oblong two-celled anthers, dorsifixed and ultimately very versatile, deeply indented at each end, and commonly exserted and pendulous. Suppression of the anterior stamen sometimes occurs (e.g.Anthoxanthum, fig. 7), or the two posterior ones may be absent (Uniola,Cinna,Phippsia,Festuca bromoides). There is in some genera (Oryza, mostBambuseae) another row of three stamens, making six in all (fig. 13, 3); andAnomochloaandTetrarrhenapossess four. The stamens become numerous (ten to forty) in the male flowers of a few monoecious genera (Pariana,Luziola). InOchlandrathey vary from seven to thirty, and inGigantochloathey are monadelphous.
Gynoecium.—The pistil consists of a single carpel, opposite the pale in the median plane of the spikelet. The ovary is small, rounded to elliptical, and one-celled, and contains a single slightly bent ovule sessile on the ventral suture (that is, springing from the back of the ovary); the micropyle points downwards. It bears usually two lateral styles which are quite distinct or connate at the base, sometimes for a greater length (fig. 14, 1), each ends in a densely hairy or feathery stigma (fig. 14). Occasionally there is but a single style, as inNardus(fig. 14, 7), which corresponds to the midrib of the carpel. The very long and apparently simple stigma of maize arises from the union of two. Many of the bamboos have a third, anterior, style.
Comparing the flower of Gramineae with the general monocotyledonous plan as represented by Liliaceae and other families (fig. 15), it will be seen to differ in the absence of the outer row and the posterior member of the inner row of the perianth-leaves, of the whole inner row of stamens, and of the two lateral carpels, whilst the remaining members of the perianth are in a rudimentary condition. But each or any of the usually missing organs are to be found normally in different genera, or as occasional developments.
1, Actual condition;
2, Theoretical, with the suppressed organs supplied.
a, Axis.
b, Flowering glume.
c, Palea.
d, Outer row of perianth leaves.
e, Inner row.
f, Outer row of stamens.
g, Inner row.
h, Pistil.
Pollination.—Grasses are generally wind-pollinated, though self-fertilization sometimes occurs. A few species, as we have seen, are monoecious or dioecious, while many are polygamous (having unisexual as well as bisexual flowers as in many members of the tribesAndropogoneae, fig. 18, andPaniceae), and in these the male flower of a spikelet always blooms later than the hermaphrodite, so that its pollen can only effect cross-fertilization upon other spikelets in the same or another plant. Of those with only bisexual flowers, many are strongly protogynous (the stigmas protruding before the anthers are ripe), such asAlopecurusandAnthoxanthum(fig. 7), but generally the anthers protrude first and discharge the greater part of their pollen before the stigmas appear. The filaments elongate rapidly at flowering-time, and the lightly versatile anthers empty an abundance of finely granular smooth pollen through a longitudinal slit. Some flowers, such as rye, have lost the power of effective self-fertilization, but in most cases both forms, self- and cross-fertilization, seem to be possible. Thus the species of wheat are usually self-fertilized, but cross-fertilization is possible since the glumes are open above, the stigmas project laterally, and the anthers empty only about one-third of their pollen in their own flower and the rest into the air. In some cultivated races of barley, cross-fertilization is precluded, as the flowers never open. Reference has already been made to cleistogamic species which occur in several genera.
Fruit and Seed.—The ovary ripens into a usually small ovoid or rounded fruit, which is entirely occupied by the single large seed, from which it is not to be distinguished, the thin pericarp being completely united to its surface. To this peculiar fruit the termcaryopsishas been applied (more familiarly “grain”); it is commonly furrowed longitudinally down one side (usually the inner, but inCoixand its allies, the outer), and an additional covering is not unfrequently provided by the adherence of the persistent palea, or even also of the floweringglume (“chaff” of cereals). From this type are a few deviations; thus inSporobolus, &c. (fig. 16), the pericarp is not united with the seed but is quite distinct, dehisces, and allows the loose seed to escape. Sometimes the pericarp is membranous, sometimes hard, forming a nut, as in some genera ofBambuseae, while in otherBambuseaeit becomes thick and fleshy, forming a berry often as large as an apple. InMelocannathe berry forms an edible fruit 3 or 4 in. long, with a pointed beak of 2 in. more; it is indehiscent, and the small seed germinates whilst the fruit is still attached to the tree, putting out a tuft of roots and a shoot, and not falling till the latter is 6 in. long. The position of the embryo is plainly visible on the front side at the base of the grain. On the other, posterior, side of the grain is a more or less evident, sometimes punctiform, sometimes elongated or linear mark, the hilum, the place where the ovule was fastened to the wall of the ovary. The form of the hilum is constant throughout a genus, and sometimes also in whole tribes.
The testa is thin and membranous but occasionally coloured, and the embryo small, the great bulk of the seed being occupied by the hard farinaceous endosperm (albumen) on which the nutritive value of the grain depends. The outermost layer of endosperm, the aleuron-layer, consists of regular cells filled with small proteid granules; the rest is made up of large polygonal cells containing numerous starch-grains in a matrix of proteid which may be continuous (horny endosperm) or granular (mealy endosperm). The embryo presents many points of interest. Its position is remarkable, closely applied to the surface of the endosperm at the base of its outer side. This character is absolute for the whole order, and effectually separates Gramineae from Cyperaceae. The part in contact with the endosperm is plate-like, and is known as thescutellum; the surface in contact with the endosperm forms an absorptive epithelium. In some grasses there is a small scale-like appendage opposite the scutellum, theepiblast. There is some difference of opinion as to which structure or structures represent the cotyledon. Three must be considered: (1) the scutellum, connected by vascular tissue with the vascular cylinder of the main axis of the embryo which it more or less envelops; it never leaves the seed, serving merely to prepare and absorb the food-stuff in the endosperm; (2) the cellular outgrowth of the axis, the epiblast, small and inconspicuous as in wheat, or larger as inStipa; (3) the pileole or germ-sheath, arising on the same side of the axis and above the scutellum, enveloping the plumule in the seed and appearing above ground as a generally colourless sheath from the apex of which the plumule ultimately breaks (fig. 17, 4,b). The development of these structures (which was investigated by van Tieghem), especially in relation to the origin of the vascular bundles which supply them, favours the view that the scutellum and pileole are highly differentiated parts of a single cotyledon, and this view is in accord with a comparative study of the seedling of grasses and of other monocotyledons. The epiblast has been regarded as representing a second cotyledon, but this is a very doubtful interpretation.
Germination.—In germination the coleorhiza lengthens, ruptures the pericarp, and fixes the grain to the ground by developing numerous hairs. The radicle then breaks through the coleorhiza, as do also the secondary rootlets where, as in the case of many cereals, these have been formed in the embryo (fig. 17, 4). The germ-sheath grows vertically upwards, its stiff apex pushing through the soil, while the plumule is hidden in its hollow interior. Finally the plumule escapes, its leaves successively breaking through at the tip of the germ-sheath. The scutellum meanwhile feeds the developing embryo from the endosperm. The growth of the primary root is limited; sooner or later adventitious roots develop from the axis above the radicle which they ultimately exceed in growth.
Means of Distribution.—Various methods of scattering the grain have been adopted, in which parts of the spikelet or inflorescence are concerned. Short spikes may fall from the culm as a whole; or the axis of a spike or raceme is jointed so that one spikelet falls with each joint as in manyAndropogoneaeandHordeae. In many-flowered spikelets the rachilla is often jointed and breaks into as many pieces as there are fruits, each piece bearing a glume and pale. One-flowered spikelets may fall as a whole (as in the tribesPaniceaeandAndropogoneae), or the axis is jointed above the barren glumes so that only the flowering glume and pale fall with the fruit. These arrangements are, with few exceptions, lacking in cultivated cereals though present in their wild forms, so far as these are known. Such arrangements are disadvantageous for the complete gathering of the fruit, and therefore varieties in which they are not present would be preferred for cultivation. The persistent bracts (glume and pale) afford an additional protection to the fruit; they protect the embryo, which is near the surface, from too rapid wetting and, when once soaked, from drying up again. They also decrease the specific gravity, so that the grain is more readily carried by the wind, especially when, as inBriza, the glume has a large surface compared with the size of the grain, or when, as inHolcus, empty glumes also take part; in Canary grass (Phalaris) the large empty glumes bear a membranous wing on the keel. In the sugar-cane (Saccharum) and several allied genera the separating joints of the axis bear long hairs below the spikelets; in others, as inArundo(a reed-grass), the flowering glumes are enveloped in long hairs. The awn which is frequently borne on the flowering glume is also a very efficient means of distribution, catching into fur of animals or plumage of birds, or as often inStipa(fig. 8) forming a long feather for wind-carriage. InTragusthe glumes bear numerous short hooked bristles. The fleshy berries of someBambuseaefavour distribution by animals.
The awn is also of use in burying the fruit in the soil. Thus inStipa, species ofAvena,Heteropogonand others the base of the glume forms a sharp point which will easily penetrate the ground; above the point are short stiff upwardly pointing hairs which oppose its withdrawal. The long awn, which is bent and closely twisted below the bend, acts as a driving organ; it is very hygroscopic, the coils untwisting when damp and twisting up when dry. The repeated twisting and untwisting, especially when the upper part of the awn has become fixed in the earth or caught in surrounding vegetation, drives the point deeper and deeper into the ground. Such grasses often cause harm to sheep by catching in the wool and boring through the skin.
A peculiar method of distribution occurs in some alpine and arctic grasses, which grow under conditions where ripening of the fruit is often uncertain. The entire spikelet, or single flowers, are transformed into small-leaved shoots which fall from the axes and readily root in the ground. Some species, such asPoa stricta, are known only in this viviparous condition; others, like our British speciesFestuca ovinaandPoa alpina, become viviparous under the special climatic conditions.
II. Classification.—Gramineae are sharply defined from all other plants, and there are no genera as to which it is possible to feel a doubt whether they should be referred to it or not. The only family closely allied is Cyperaceae, and the points of difference between the two may be here brought together. Thebest distinctions are found in the position of the embryo in relation to the endosperm—lateral in grasses, basal in Cyperaceae—and in the possession by Gramineae of the 2-nerved palea below each flower. Less absolute characters, but generally trustworthy and more easily observed, are the feathery stigmas, the always distichous arrangement of the glumes, the usual absence of more general bracts in the inflorescence, the split leaf-sheaths, and the hollow, cylindrical, jointed culms—some or all of which are wanting in all Cyperaceae. The same characters will distinguish grasses from the other glumiferous orders, Restiaceae, and Eriocaulonaceae, which are besides further removed by their capsular fruit and pendulous ovules. To other monocotyledonous families the resemblances are merely of adaptive or vegetative characters. Some Commelinaceae and Marantaceae approach grasses in foliage; the leaves ofAllium, &c., possess a ligule; the habit of some palms reminds one of the bamboos; and Juncaceae and a few Liliaceae possess an inconspicuous scarious perianth. There are about 300 genera containing about 3500 well-defined species.
The great uniformity among the very numerous species of this vast family renders itsclassificationvery difficult. The difficulty has been increased by the confusion resulting from the multiplication of genera founded on slight characters, and from the description (in consequence of their wide distribution) of identical plants under several different genera.
No characters for main divisions can be obtained from the flower proper or fruit (with the exception of the character of the hilum), and it has therefore been found necessary to trust to characters derived from the usually less important inflorescence and bracts.
Robert Brown suggested two primary divisions—Paniceae and Poaceae, according to the position of the most perfect flower in the spikelet; this is the upper (apparently) terminal one in the first, whilst in the second it occupies the lower position, the more imperfect ones (if any) being above it. Munro supplemented this by another character easier of verification, and of even greater constancy, in the articulation of the pedicel in the Paniceae immediately below the glumes; whilst in Poaceae this does not occur, but the axis of the spikelet frequently articulatesabovethe pair of empty basal glumes. Neither of these great divisions will well accommodate certain genera allied toPhalaris, for which Brown proposed tentatively a third group (since namedPhalarideae); this, or at least the greater part of it, is placed by Bentham under the Poaceae.
The following arrangement has been proposed by Professor Eduard Hackel in his recent monograph on the order.