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species—crosspollination (or allogamy). Effective pollination may also occur between flowers of different species, or occasionally, as in the See also:

case of several See also:

ORCHIDS

orchids, of different genera—this is known as hybridization. The method of pollination is to some extent governed by the See also:

DISTRIBUTION (Lat. distribuere, to deal out)

distribution of the stamens and pistil. In the case of unisexual flowers, whether monoecious, that is, with staminate and pistillate flowers on one and the same plant, such as many of our native trees—See also:

OAK (O. Eng., (lc)

oak, See also:

BEECH

beech, See also:

birch, See also:

ALDER

alder, &c., or dioecious with staminate and pistillate flowers on different See also:

PLANTS

plants, as in willows and poplars, See also:

CROSS

cross pollination only is possible. In bisexual or hermaphrodite flowers, that is, those in which both stamens and pistil are See also:

PRESENT

present, though self-pollination might seem the obvious course, this is often prevented or hindered by various arrangements which favour cross-pollination. Thus the anthers and stigmas in any given flower are often mature at different times; this See also:

CONDITION (Lat. condicio, from condicere, to agree upon, arrange; not connected with conditio, from condere, conditum, to put together)

condition, which is known as dichogamy and was firstpointed out by See also:

SPRENGEL, KURT (1766-1833)

Sprengel, may be so well marked that the stigma has ceased to be receptive before the anthers open, or the anthers have withered before the stigma becomes receptive, when cross-pollination only is possible, or the stages of maturity in the two See also:

ORGANS

organs are not so distinct, when self-pollination becomes possible later on. The flower is termed proterandrous or proterogynous according as anthers or stigmas mature first. The See also:

TERM

term homogamy is applied to the simultaneous maturity of stigma and anthers. Spontaneous ' self-pollination is rendered impossible in some homogamous flowers in consequence of the relative position of the anthers and stigma—this condition has been termed herkogamy. Flowers in which the relative position of the organs allows of spontaneous self-pollination may be all alike as regards length of See also:

style and stamens (homomorphy or homostyly), or differ in this respect (heteromorphy) the styles (From Strasburger's Lehrbuch der Botanik, by permission of Gustav See also:

Fischer.) G, Level of stigma; S, level of anthers; P, N, pollen grains and stigmatic papillae of See also:

long-styled form; p, n, See also:

DITTO (from the Lat. dictum, something said, Ital. detto, aforesaid)

ditto of See also:

SHORT, FRANCIS JOB (1857– )

short-styled form.

and stamens being of different lengths in different flowers (heterostyly) or the stamens only are of different lengths (heteranthery). Flowers which are closed at the See also:

time of maturity of anthers and stigmas are termed cleistogamous. Self-pollination is effected in very various ways. In the simplest case the anthers are See also:

close to the stigmas, covering these with pollen when they open; this occurs in a number of small See also:

ANNUAL

annual plants, also in See also:

NARCISSUS

Narcissus, See also:

CROCUS

Crocus, &c. In See also:

SNOWDROP

snowdrop and other pendulous flowers the anthers form a See also:

CONE (Gr. Kwvos)

cone around the style and the pollen falls on to the underlying stigmas, or in erect flowers the pollen may fall on to the stigmas which See also:

lie directly beneath the opening anthers (e.g. Narthecium). In very many cases the pollen is carried to the stigma by See also:

ELONGATION

elongation, curvature or some other See also:

MOVEMENT

movement of the filament, the style or stigma, or corolla or some other See also:

PART

part of the flower, or by correlated movements of two or more parts. For instance, in many flowers the filaments are at first directed outwards so that self-pollination is not possible, but later incline towards the stigmas and pollinate them (e.g. numerous See also:

SAXIFRAGACEAE

Saxifragaceae, See also:

CRUCIFERAE

Cruciferae and others), or the style, which first projects beyond the anthers, shortens later on so that the anthers come into contact with the stigmas (e.g. species of Cactaceae), or the style bends so that the stigma is brought within the range of the pollen (e.g. species of Oenothera, Epilobium, most See also:

MALVACEAE

Malvaceae, &c.). In Mirabilis See also:

JALAPA, XALAPA, or HALAPA

Jalapa and others the filaments and style finally become intertwined, so that pollen is brought in contact with the stigma. Self-pollination frequently becomes possible towards the end of the See also:

LIFE

life of a flower which during its earlier stages has been capable only of cross-pollination. This is associated with the fact, so ably demonstrated by See also:

Darwin, that, at any See also:

RATE

rate in a large number of cases, cross-pollination yields better results, as measured by the number of seeds produced and the strength of the offspring, than self-pollination; the latter is, however, preferable to See also:

ABSENCE (Lat. absentia)

absence of pollination. In many cases pollen has no effect on the stigma of the same flower, the plants are self-sterile, in other cases See also:

EXTERNAL

external pollen is more effective (pre-potent) than pollen from the same flower; but in a very large number of cases experiment has shown that there is little or no difference between the effects of external pollen and that from the same flower.

Cross-pollination may occur between two flowers on the same plant (geitonogamy) or between flowers on distinct plants (xenogamy). The former, which is a somewhat less favourable method than the latter, is effected by See also:

air-currents, See also:

INSECT

insect agency, the actual contact between stigmas and anthers in neighbouring flowers, where, as in the See also:

FAMILY

family See also:

COMPOSITAE

Compositae, flowers are closely crowded, or by the fall of the pollen from a (From Darwin's Different Forms of Flowers by permission.) The dotted lines with the arrow show the directions in which pollen must be carried to each stigma to ensure full fertility. higher on to the stigmas of a See also:

LOWER

lower flower. Anton See also:

KERNER, JUSTINUS ANDREAS CHRISTIAN (1786–1862)

Kerner has shown that crowded inflorescences such as those of Compositae and See also:

UMBELLIFERAE

Umbelliferae are especially adapted for geitonogamy. Xenogamy is of course the only possible method in diclinous plants; it is also the usual method in monoclinous plants, owing to the fact that stamens and carpels often mature at different times (dichogamy), the plants being proterandrous or proterogynous. Even in homogamous flowers cross-pollination is in a large proportion of cases the effective method, at any rate at 'first. owing to the relative position of anther and stigma or the fact that the plant is self-sterile. The subject of heterostyly was investigated by Darwin (see his Forms of Flowers) and later by See also:

HILDEBRAND, LAY OF (Deis Hildebrandslied)

Hildebrand. In the case of a dimorphic flower, such as Primula, four modes of pollination are possible, two distinguished by Darwin as legitimate, between anthers and stigmas on corresponding levels, and two so-called illegitimate unions, between anthers and stigmas at different levels (cf. fig. I). In a trimorphic flower such as Lythrum salicaria there are six possible legitimate unions and twelve illegitimate (see fig. 2). Experiment showed that legitimate unions yield a larger quantity of See also:

SEED (from the root seen in Lat. serere, to sow)

seed than illegitimate.

Many plants produce, in addition to See also:

ORDINARY (med. Lat. ordinarius, Fr. ordinaire)

ordinary open flowers, so-called cleistogamous flowers, which remain permanently closed but which notwithstanding produce See also:

FRUIT (through the French from the Lat. fructus; frui, to enjoy)

fruit; in these the corolla is inconspicuous or absent and the pollen grows from the anther on to the stigma of the same flower. Species of See also:

VIOLA [Fr. viole, Ger. Bratsche, Ital. viola, alto]

Viola (see fig. 3), See also:

OXALIS

Oxalis acetosella (See also:

wood See also:

SORREL

sorrel) and Lamium amplexicaule are commonly occurring in-stances. The cleistogamous flowers are See also:

DEVELOPED

developed before or after the normal open flowers at seasons less favourable for cross-pollination. In some cases flowers, which open under normal circumstances, remain closed owing to unfavourable circumstances, and self-pollination occurs as in a FIG.3.—Cleistogamous typical cleistogamous flower--these flower of Viola sylvatica. have been distinguished as pseudo- r, flower X 4. cleistogamous. Instances occur in flower mou hily magnified and c cut oghpen. See also:

WATER

water plants, where flowers are un- a, anther; s, pistil; able to reach the surface (e.g. Alisma st, style; v, stigmatic natans, water See also:

BUTTERCUP

buttercup, &c.) or surface. where flowers remain closed in dull or See also:

COLD (in O. Eng. cald and ceald, a word coming ultimately from a root cognate with the Lat. gelu, gelidus, and common in the Teutonic languages, which usually have two distinct forms for the substantive and the adjective, cf. Ger. Kolte, kalt, Dutch koude

cold See also:

WEATHER (O. Eng. weder; the word is common to Teutonic languages; cf. Du. weder, Dan. veir, Icel. ve8r, and Ger. Wetter and Gewitter, storm; the root is wa- to blow, from which is derived " wind ")

weather. Systems of See also:

CLASSIFICATION (Lat. classis, a class, probably from the root cal-, cla-, as in Gr. icaMce, clamor)

classification of flowers according to the agency by which pollination is effected have been proposed by Delpino, H. See also:

Muller and other workers on the subject.

Knuth suggests the following, which is a modification of the systems proposed by Delpino and Muller. A. Water-pollinated plants, Hydrophilae. A small See also:

GROUP

group which is subdivided thus: a. Pollinated under the water; e.g. Najas where the pollen grains are rather heavier than water, and sinking down are caught by the stigmas of the extremely See also:

SIMPLE

simple See also:

FEMALE

female flowers. b. Pollination on the surface, a more frequent occurrence than (a). In these the pollen floats on the surface and reaches the stigmas of the female flowers as in Callitriche, Ruppio, Zostera, Elodea. In Vallisneria (fig. 4) the male flowers become detached and See also:

FLOAT (in O. Eng. floc and flota, in the verbal form f eotan; the Teutonic root is flut-, another form of flu-, seen in " flow," cf. " fleet "; the root is seen in Gr. a-M e, to sail, Lat. pluere, to rain; the Lat, fluere and fluctus, wave, is not connect

float on the surface of the water; the anthers are thus brought in contact with the stigmas of the female flowers. B.

Wind-pollinated plants, Anemophilae.—In these the pollen grains are smooth and See also:

LIGHT

light so as to be easily blown about, and are produced in great quantity; the stigmas are See also:

brush like or feathery, and usually long and protruding so as readily to catch the pollen. As no means of attraction are required the flowers are inconspicuous and without scent or See also:

NECTAR

nectar. The male inflorescence is often a pendulous catkin, as in See also:

HAZEL (0. Eng. hresell; cf. Ger. Hasel, Swed. and Dan. Kassel, &c.,; Fr. noiselier, coudrier)

hazel and many native See also:

ENGLISH

English trees (fig. 5) ; or the anthers are loosely fixed on long See also:

THREAD (0. Eng. praed, literally, that which is twisted, prawan, to twist, to throw, cf. " throwster," a silk-winder, Ger. drehen, to twist, turn, Du. draad, Ger. Draht, thread, wire)

thread-like filaments as in See also:

GRASSES

grasses (fig. 6). A, female flower; s, stigmas. B, male flowers; I before; 2, after spreading of the petals. A male flower has floated alongside a female and one of its anthers, which have opened to set See also:

FREE

free the pollen, is in contact with a stigma. a, anther. C. See also:

Animal-pollinated plants, Zoidiophilae, are subdivided according to the See also:

KIND (O. E. ge-cynde, from the same root as is seen in " kin," supra)

kind of animal by agency of which pollination is effected, thus: a. See also:

Bat-pollinated, Chiropterophilae.—A Freycinetia, native of See also:

JAVA

Java, and a species of Bauhinia in See also:

TRINIDAD

Trinidad are visited by bats which See also:

TRANSFER (from Lat. transferre, to bear across, carry over)

transfer the pollen. Long. stq led.

See also:

Mid. styled Short-styled. b. See also:

BIRD

Bird-pollinated, Ornithophilae.—Humming-birds and See also:

HONEY (Chin. me ; Sansk. madhu, mead, honey; cf. A.S. medo, medu, mead; Gr. p Xt, in which 0 or b is changed into X; Lat. mel ; Fr. miel ; A.S. hung ; Ger. Honig)

honey-suckers are agents of pollination in certain tropical plants; they visit the generally large and brightly-coloured flowers either for the honey which is secreted in considerable quantity or for the insects which have been attracted by the honey (fig. 7). Flowers of Hazel. truding hairy stigmas. c. See also:

SNAIL

Snail or slug-pollinated flowers, Malacophilae.—In small flowers which are crowded at the same level or in See also:

FLAT (a modification of O. Eng. flet, an obsolete word of Teutonic origin, meaning the ground beneath the feet)

flat flowers in which the stigmas and anthers project but little, slugs or snails creeping over their surface may transfer to the stigma the pollen which clings to the slimy See also:

FOOT

foot. Such a transfer has been described in various Aroids, Rohdea japonica (See also:

LILIACEAE

Liliaceae), and other plants. (From a See also:

DRAWING

drawing in the Botanical See also:

GALLERY (through Ital. galleria, from Med. Lat. galeria, of which the origin is unknown)

Gallery at the See also:

BRITISH

British Museum.) d. Insect-pollinated, Entomophilae, a very large class characterized by sticky pollen grains, the surface of which bears spines, warts or other projections (fig. 8) which facilitate See also:

ADHESION (from Lat. adhaerere, to adhere)

adhesion to some part of the insect's See also:

BODY

body, and a relatively small stigma with a sticky surface. The flowers have an attractive floral envelope, are scented and often contain honey or a large amount of pollen: by these means the insect is enticed to visit it.

The form, colour and scent of the flower vary widely, according to the class of insect whose aid is sought, and there are also numerous devices for protecting the pollen and nectar from See also:

RAIN (O.E. regn; the word is common to Teutonic languages, cf. Ger. Regen, Swed. and Dan. regn; it has been connected with Lat. rigare, to wet, Gr. 13pixeav)

rain and See also:

dew or from the visits of those insects which would not serve the purpose of pollen-transference (unbidden guests).' The following subdivisions have been suggested A. Pollen Flowers.—These offer only pollen to their visitors, as species of See also:

ANEMONE

anemone, See also:

POPPY

poppy, See also:

rose, See also:

TULIP (Tulipa)

tulip, &c. They are simple in structure and See also:

REGULAR

regular in form, and the generally abundant pollen is usually freely exposed. B. Nectar Flowers.—These contain nectar and include the following See also:

groups: i. Flowers with exposed nectar, readily visible and accessible to all visitors. These are very simple, open and generally regular flowers, See also:

white, greenish-yellow or yellow in colour and are chiefly visited by insects with a short See also:

PROBOSCIS

proboscis, such as short-tongued wasps and flies, also beetles and more rarely bees. Examples are Umbelliferae as a family, saxifrages, See also:

HOLLY (Ilex Aquifolium)

holly, Acer, Rhamnus, See also:

EUONYMUS

Euonymus, See also:

EUPHORBIA

Euphorbia, &c. 2. Flowers with nectar partly concealed and visible only in See also:

bright See also:

SUNSHINE

sunshine. The generally regular flowers are completely open only in bright sunshine, closing up into cups at other times. Such are most Cruciferae, buttercups, See also:

king-See also:

CUP (in O.E. cuppe; generally taken to be from Late Lat. cuppa, a variant of Lat. cupa, a cask, cf. Gr. KuaeXXov)

cup (Caltha), See also:

POTENTILLA (nat. order Rosaceae, q.v.)

Potentilla.

White and yellow See also:

COLOURS, MILITARY

colours predominate and insects with a proboscis of See also:

MEDIUM

medium length are the See also:

COMMON

common pollinating agents, such as short-tongued bees. 3. Flowers with nectar concealed by pouches, hairs, &c. Regular flowers predominate, e.g. See also:

GERANIUM

Geranium, Cardamine pratensis, mallows, Rubus, Oxalis, Epilobium, &c., but many species show more or less well-marked median symmetry (zygomorphism) as Euphrasia, Orchis, See also:

THYME

thyme, &c., and red, See also:

BLUE (common in different forms to most European languages)

blue and See also:

VIOLET

violet are the usual colours. Long-tongued insects such as the honey-See also:

BEE (Sanskrit blza, AS. ben, Lat. apis)

bee are the most frequent visitors. 4. Social flowers, whose nectar is concealed as in (3), but the flowers are grouped in heads which render them strikingly conspicuous, and several flowers can be simultaneously pollinated. Such are Compositae as a class, also Scabiosa, Armeria (See also:

sea-See also:

PINK

pink) and others. 5. Hymenopterid flowers, which fall into the following groups: Bee-flowers proper, humble-bee flowers requiring a longer proboscis to reach the nectar, See also:

WASP (Lat. vespa)

wasp-flowers such as fig-wort (Scrophularia nodosa) and See also:

ICHNEUMON (Gr. ixveu)

ichneumon flowers such as tway-blade (Listera ovata). The shapes and colours are extremely varied; bilaterally symmetrical forms are most frequent with red, blue or violet colours.

Such are Papilionaceous flowers, Violaceae, many See also:

LABIATAE (i.e. " lipped," Lat. labium, lip)

Labiatae, See also:

SCROPHULARIACEAE

Scrophulariaceae and others. Many are highly specialized so that pollination can be effected by a few species only. Examples of more See also:

SPECIAL

special mechanisms are illustrated by See also:

SALVIA

Salvia (fig. 9). The long connective of the single stamen is hinged to the short filament and has a shorter See also:

ARM (a common Teutonic word; the Indo-European root is ar, to join or fit; cf. the Lat. armus, shoulder, and the plural word arma, weapons, Gr. apphs, joint, and the reduplicated apapilKSV, to join)

arm ending in a See also:

blunt See also:

PROCESS

process and a longer arm bearing a See also:

HALF

half-anther. A large bee in probing for honey comes in contact with the end of the short arm of the See also:

lever and causes the longer arm to descend and the pollen is deposited on the back of the insect (fig. 9, I). In a later See also:

STAGE (Fr. 6/age; from Lat. stare, to stand)

stage (fig. 9, 2) the style elongates and the forked stigma occupies the same position as the anther in fig. 9, I. f, Flower visited by a humble-bee, showing the See also:

PROJECTION

projection of the curved connective bearing the anther from the See also:

HELMET (from an obsolete diminutive of O. Fr. helme, mod. heaume; the English word is " helm," as in O. Eng., Dutch and Ger.; all are from the Teutonic base hal-, pre-Teut. kal-, to cover; cf. Lat. celare, to hide, Eng. " hell," &c.)

helmet-shaped upper See also:

LIP (a word common in various forms, to Teutonic languages, cf Ger. Lippe, Dan. laebe; Lat. labium is cognate)

lip and the deposition of the pollen on the back of the humble-bee. 2, Older flower,with connective See also:

DRAWN

drawn back, and elongated style.

i See A. Kerner, Plants and their Unbidden Guests. 1 2 (From Strasburger's Lehrbuck der Botanik'by permission of Gustav Fischer.) 4, The staminal apparatus at See also:

REST (O. Eng. rest, reste, bed, cognate with other Teutonic forms, e.g. Ger. Rast, Riiste, rest, and probably Gothic Rasta, league, i.e. resting or stopping place)

rest, with connective enclosed within the upper lip. 3, The same, when disturbed by the entrance of the proboscis of the bee in the direction of the arrow; f, filament; c, connective; s, the obstructing half of the anther. In See also:

BROOM

Broom there is an explosive machanism ; the pressure of the insect visitor on the See also:

KEEL

keel of the corolla causes a sudden See also:

RELEASE (O.Fr. reles, variant of relais, from relaisser, to release, let go, Lat. relaxare)

release of the stamens and the scattering of a See also:

CLOUD (from the same root, if not the same word, as " clod," a word common in various forms to Teutonic languages for a mass or lump; it is first applied in the usual sense in the late 13th century; the Anglo-Saxon chid is only used in the sense of " a ma

cloud of pollen over its body. 6. Lepidopterid flowers, visited chiefly by Lepidcptera, which are able to reach the nectar concealed in deep, narrow tubes or spurs by means of their long slender proboscis. Such are: (a) Butterfly-flowers, usually red in colour, as Dianthus carthusianorum; (b) See also:

MOTH

Moth-flowers, white or whitish, as See also:

HONEYSUCKLE (Mid. Eng., honysocle, i.e. any plant from which honey may be sucked,—cf. A.-S. huni-suge, privet; Ger. Geissblalt; Fr. chevrefeuille)

honeysuckle (Loniceta periclymenum). 7. See also:

FLY (formed on the root of the supposed original Tent. fleugan, to fly)

Fly flowers, chiefly visited by See also:

DIPTERA (Sir, double, arepa, wings)

Diptera, and including very different types: a. Nauseous flowers, dull and yellowish and dark See also:

PURPLE

purple in colour and often spotted, with a See also:

SMELL (connected etymologically with " smoulder " and " smoke ")

smell attractive to carrion flies and dung flies, e.g. species of Saxifraga. b.

Pitfall flowers such as Asarum, See also:

ARISTOLOCHIA (Gr. apwros, best, Xoxeia, child-birth, in allusion to its repute in promoting child-birth)

Aristolochia and Arum maculation, when the insect is caught and detained until pollination is effected (fig. Io). c. Pinch-See also:

TRAP (0. Eng. treppe or traepee, properly a step, as that or which an animal places its foot and is caught, cf. Ger. Treppe flight of stairs)

trap flowers, as in the family Asclepiadaceae, where the proboscis, claw or bristle of the insect is caught in the clip to which the pairs of pollinia are attached. Bees, wasps and larger insects serve as pollinating agents p, Pistillate, s, staminate flowers; h, sterile flowers forming a circlet of stiff hairs closing the mouth of the chamber formed by the lower part of the spathe. s, One of the scales which form the coronet in the flower, enlarged.d. Deceptive flowers such as Parnassia, where the conspicuous coronet of glistening yellow balls suggests a plentiful See also:

SUPPLY (through Fr. from Lat. supplere, to fill up)

supply of nectar drops (fig. II). e. Hoverfly flowers, small flowers which are beautifully coloured with radiating streaks pointing to a sharply-defined centre in which is the nectar, as In See also:

VERONICA, ST

Veronica chamaedrys (fig. 12). own pollen." In 1999 an Englishman, (From Vines's See also:

TEXT (Lat. textum, woven fabric, from texere, to weave)

Text See also:

BOOK

Book of See also:

Thomas See also:

Andrew See also:

Knight, after experiments Botany, by permission.) on the cross-fertilization of cultivated FIG.

12.—Flower of plants, formulated the conclusion that no Veronica. plant fertilizes itself through many genera- k, Calyx. tions. Sprengel's See also:

WORK

work, which had been u, u, u, The three lobes almost forgotten, was taken up again by of the lower lip of See also:

Charles Darwin, who concluded that no the rotate corolla. organic being can fertilize itself through o, The upper lip. , an unlimited number of generations; but s, s, The two stamens, a cross with other individuals is occasion- n, The stigma. ally—perhaps at very long intervals—indis- pensable. Darwin's See also:

WORKS

works on dimorphic flowers and the fertilization of orchids gave powerful support to this statement. The study of the fertilization, or as it is now generally called " pollination," of flowers, was continued by Darwin and taken up by other workers, notably See also:

FRIEDRICH, JOHANN (1836— )

Friedrich Hildebrand, Federico Delpino and the See also:

BROTHERS, RICHARD (1757-1824)

brothers Fritz and See also:

Hermann Muller. Hermann Muller's work on The Fertilization of Flowers by Insects and their Reciprocal Adaptations (1873), followed by subsequent works on the same lines, brought together a great number of observations on floral mechanisms and their relation to insect-visits. Muller also suggested a modification of the I{See also:

NIGHT

night-Darwin See also:

law, which had See also:

LEFT

left unexplained the numerous instances of continued successful self-pollination, and restated it on these terms: " Whenever offspring resulting from See also:

CROSSING

crossing comes into serious conflict with offspring resulting from self-fertilization, the former is victorious. Only where there is no such struggle for existence does self-fertilization often prove satisfactory for many generations." An increasing number of workers in this See also:

field of plant See also:

BIOLOGY (Gr. Qios, life)

biology in See also:

England, on the See also:

CONTINENT (from Lat. continere, "to hold together "; hence " connected," " continuous ")

Continent and in See also:

AMERICA

America has produced a great See also:

mass of observations, which have recently been brought together in Dr See also:

Paul Knuth's classic work, Handbook of Flower Pollination, an English See also:

TRANSLATION

translation of which has been published (1908) by the See also:

Clarendon See also:

PRESS (through Fr. presse from Lat. pressare, frequentative of premere, to crush, squeeze, press)

Press.

End of Article: POLLINATION

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