FOOTNOTES:

Leafy.Floral.Ranunculaceæ.Clematis.Anemone!*Anemone!Ranunculus!*Ranunculus!Delphinium.Caltha.Aquilegia!Cruciferæ.Bunias.*Cheiranthus!Erucago.*Matthiola!Sisymbrium!Brassica!Nasturtium.Hesperis.Sinapis!Diplotaxis.Lunaria.Erysimum.Alyssum.Peltaria.Cardamine!Cleome.Cistaceæ.Helianthemum!Caryophylleæ.Dianthus!*Dianthus!Silene!Lychnis!Violaceæ.Viola!Tiliaceæ.Triumfetta!Geraniaceæ.Geranium!Sapindaceæ.Pavia!Pavia!Malvaceæ.Paritium.Hibiscus!Malpighiaceæ.Byrsonima!Rutaceæ.Genera not specified.*Dictamnus!Resedaceæ.Reseda.Caylussa!Aurantiaceæ.*Citrus!Vitaceæ.Vitis.Vitis.Umbelliferæ.Heracleum.Angelica.Thysselinum.*Athamanta.*Daucus!*Torilis.Rosaceæ.*Rosa!*Rosa!*Geum!*Geum!Agrimonia.Amygdalus.Prunus!Spiræa!Spiræa!Rubus.*Pyrus!*Pyrus!?LeguminosæTrifolium!Medicago!Melilotus.Pisum!Cucurbitaceæ.Cucumis.Passifloraceæ.Passiflora.Philadelphaceæ.Philadelphus.Onagraceæ.Epilobium!Epacridaceæ.Epacris!Ericaceæ.*Erica.Rhododendron!Convolvulaceæ.Convolvulus.Gentianaceæ.Gentiana.Gentiana.Apocynaceæ.Vinca.Jasminaceæ.Jasminum!Scrophulariaceæ.Verbascum!Antirrhinum!*Digitalis!*Linaria!Veronica.Orobanchaceæ.Orobanche.Labiatæ.Genera not specified.Stachys.Phlomis!Hydrophyllaceæ.Hydrophyllum.Boraginaceæ.Anchusa.Symphytum.Primulaceæ.*Dodecatheon.*Cortusa.*Anagallis!*Anagallis!*Primula.Dipsacaceæ.Scabiosa.Compositæ.Hieracium!Hieracium!Cirsium.Cirsium.Hypochæris.Calendula!Spilanthes.Carthamus.Coreopsis.Campanulaceæ.Campanula.*Campanula!Polygonaceæ.Genera not specified.Rumex.Santalaceæ.Thesium.Liliaceæ.Genera not specified.Tulipa!Hemerocallis!Asphodelus.Hyacinthus!Iridaceæ.Iris.Amaryllidaceæ.Narcissus!Leucojum.Orchidaceæ.Orchis!Habenaria.Cyperaceæ.Carex.Gramineæ.Phleum.

Axillary prolificationis the term applied to those cases wherein one or more adventitious buds spring from the axils of one or more of the parts of the flower. Engelmann makes use of the word ecblastesis to denote the same condition. Both terms are open to the objection that they do not clearly enable us to distinguish prolification occurring within the flower from a similar state originating outside the flower, within the bracts of the inflorescence. This latter condition, called by Moquin-Tandon lateral prolification (see Prolification of the Inflorescence), is as truly axillary as that to which the name is restricted. In consequence of certain peculiarities in the structure of some flowers, to be hereafter alluded to, it is not in all cases easy to decide whether the new growth springs from theinterior of the flower, or from the inflorescence beneath the flower.

The accessory bud presents itself as a leaf-bud, a branch, a flower-bud, or a miniature inflorescence; it may be sessile, but is far more frequently stalked, and in more than half the number of cases it is a flower-bud or an inflorescence. There may be one or more of these buds; if two only, then they are usually placed directly opposite one to the other, on the opposite sides of the flower.

It will be seen, from the appended list, that the orders and genera in which this description of adventitious growth occurs most frequently are the following:—Cruciferæ, especially the genusBrassica;Caryophyllaceæ, e.g.Dianthus;Resedaceæ;Leguminosæ, e.g.Melilotus,Trifolium, &c.;Rosaceæ, e.g.Rosa,Potentilla, &c.;Umbelliferæ, andCampanulaceæ. For the most part, these are groups also peculiarly liable to central prolification.

All the parts of the flower may be thus affected; but, as might have been anticipated from the foliaceous nature of the sepals, the new bud usually arises from within the axil of one of those organs. Next in frequency to the calyx, the pistil is subjected to this change—the carpels in such a case being disunited and leaf-like. The petals rank next, and lastly the stamens; these latter, indeed, are usually, but not invariably, absent, the new growth occupying their position. Hence it may well be that when such is the case, there is no real axillary prolification, but rather the substitution of a bud for a stamen. Generally, however, the position of the accessory bud is such that it may properly be referred to the axil of an undeveloped or rudimentary stamen.

The largest number of instances of this malformation, not merely generically, but also individually, occurs in plants the members of whose floral whorls are not united one to the other; thus, it is far more common in polypetalous plants than in gamopetalousones. In the prolified flowers belonging to the latter group, the sepals, if not actually uncombined, are only united for a short distance. The same relationship, but in a much less degree, exists in the case of median prolification, as that aberration is likewise most commonly met with in polypetalous flowers. Another feature of interest is the rarity with which axillary prolification is found in irregular gamopetalous blooms. It may be that the irregular and comparatively excessive growth in some parts of these flowers, as compared with others, may operate in checking any luxuriant tendency in other directions.

As in the case of median prolification, plants having an indefinite inflorescence are more liable to be affected with ecblastesis than those having a definite one. The degree of branching of the inflorescence may be noticed, as this deformity is far more common in plants whose peduncles are branched than in those which have either a solitary flower or an unbranched flower-stalk. More than two thirds of the entire number of genera cited as the subjects of this malformation have a branched inflorescence of some form or other; and about two thirds of the cases occur in genera having some form of indefinite inflorescence. If individual instances could be accurately computed, the proportion would be even higher.

Fully three fourths of the entire number of genera recorded as occasionally the subjects of this irregularity possess in their usual state some peculiarity of the thalamus; for instance, in about a third of the whole number of genera the thalamus is more or less prolonged between some or other of the floral whorl, e.g.Caryophyllaceæ,Potentilla,Anemone,Dictamnus,Umbelliferæ, &c. About one fourth of the genera have numerous stamens or numerous carpels, or both, springing naturally from the thalamus. In others (about one sixth) the thalamus is enlarged into a disc, or else presents one or moreglandular swellings,e.g.Reseda,Nymphæa,Cruciferæ. In the last-named family, as has been already remarked, prolification is very common. It would be interesting to ascertain precisely what part of an inflorescence is most liable to this affection; but as information on this point is but rarely given in the records of these cases, I can only give the results of my own observations, which go to show that, in a many-flowered inflorescence, those flowers at the outside, or at the lower portion, seem to be more frequently the subjects of this change than those situated elsewhere. This may probably be accounted for by the fact that the malformation is met with most generally in plants with an indefinite form of inflorescence, and therefore the lowermost or outermost flowers are most fully nourished; the upper flowers being in a less advanced condition, the change is more likely to be overlooked in them; or it may be that from the unusual luxuriance in the lower flowers, the upper ones may be either present in their ordinary condition, or may be (as indeed frequently happens) stunted in the size and proportion of their several parts.

Axillary foliar prolification of the flower.—The formation of an adventitious leaf-bud in the axil of any of the parts of the flower is not of such common occurrence as the development of a flower-bud in similar situations, nor is it so frequent as median foliar prolification. I have seen leafy shoots proceeding from the axils of the sepals in the flowers ofBrassica, and a similar occurrence has been noticed inCaltha palustris,Herreria parviflora, and other plants. Dr. Marchand's flowers ofAnagallis, previously referred to at p. 117, showed good illustrations of this occurrence, as also some specimens described by Kirschleger inA. phœnicea.[134]Steinheil has figured and described[135]a flower ofScabiosain which there was an adventitious formation of leafy shoots in the axil of the outer calyx. In some flowers, such asConvolvulus,Anemone, &c., the exact nature of the sub-floral leaves is uncertain,i.e.it is open to doubt whether the organs in question are bracts or leaves pertaining to the inflorescence, or whether they are really parts of the flower. When leafy shoots are formed in the axils of such organs, the adventitious growth may be referred to extra-floral prolification, prolification of the inflorescence that is, or to axillary prolification, according to the view taken of the real nature of the sub-floral leaves. So far as the mere occurrence of prolification is concerned, it is not very material which view be adopted. The same remark applies to cases where leaf-buds occur on the outer surface of inferior ovaries, as inRosaceæ,Pomaceæ,Philadelphus, orTetragonia expansa, as elsewhere mentioned.

It would seem more consistent with the general arrangements of parts, that the adventitious buds should be formed more frequently outside than within the flower proper.

Knight[136]figures and describes the occurrence of small tubers or fleshy leaf-buds in the axils of the sepals of a potato, a curious illustration of the real morphological nature of the tuber.

Axillary floral prolification of the flower.—As already stated, this is of more common occurrence than the formation of a leaf-bud in a similar situation. Any of the parts of the flower may thus subtend a flower-bud, though probably the new buds more frequently originate in the axils of the sepals than in the other whorls. InCruciferæthe change in question is, relatively speaking, very common. In cauliflowers and broccoli I have frequently met with stalked flowers proceeding from the axils of the sepals, so also in some fuchsias I have seen a ring of stalked flower-buds alternatingwith the petals, which, together with the stamens and pistil, remained unaffected. The number of parts in the supernumerary structures is generally less than the normal flowers.

In Mr. Herbert Spencer's 'Principles of Biology,' part iv, p. 37, are figured and described some monstrous inflorescences inAngelicaand otherUmbelliferæ, from which, amongst other things, the author draws the conclusion that there is no absolute distinction between leaf and branch. Without staying for the moment to discuss this matter, it may here be said that the Umbellifers in question apparently owe their peculiarities rather to axillary prolification within the flower, or to prolification of the inflorescence, than to an actual transformation of a flower or any portion of a flower into an umbellule.[137]

In the 'Gardeners' Chronicle,' 1855, p. 551, an instance is figured of the production of a supernumerary flower proceeding from the axil of a stamen in a species ofNymphæa(fig. 65). The ovary in this case was wanting, but in its place was a tuft of small leaves. It is curious that among Dr. Kirk's drawings of east tropical African plants now at Kew, there should be one representing a precisely similar state of things. The species in both instances wasNymphæa Lotus, or a cultivated variety of it.

M. Wesmael[138]describes a very singular case of what appears to have been referable to axillary prolification in the flowers ofCarex acuta. The rachillus is described as prolonged through the utricle by the side of the stigmas, bearing on its side a bract, then a secondary utricle, from the axil of which sprung a short stem surmounted by an ovary. Wigand, 'Flora,' 1856, mentions a similar change inCarex glauca. In this instance the base of the female inflorescence bore lateral spikes, which projected from the utricles; some of these adventitious spikes were female, othersfemale below and male above, others, again, wholly male.

Fig.65.—Flower ofNymphæa Lotus, var., showing axillary floral prolification. The section also shows the tuft of leaves that occupied the place of the ovary.

Various changes in the form and arrangement of the several floral whorls accompany axillary prolification; some of these affect the particular organ or organs implicated, and these only, while in other cases some other parts of the flower likewise undergo modification. The changes most commonly met with are such as may be classed under Goethe's theory of retrograde metamorphosis; for instance, if a supplementary bud be developed in the axil of a sepal, that sepal is likely to be more than ordinarily leaf-like in appearance. The dislocation of the affected sepal from its fellows is a very frequent occurrence; in cases of thiskind the detached sepal is placed below the others, thus approximating, in position as well as in function, to the bracts. In some of the instances of proliferous pears, on which I shall have occasion to comment, the sepals are described as sharing in the succulent character of the fruit.

The petals, under such circumstances, often exist in the guise of sepals or of small leaves; and instances are recorded wherein the place of the calyx and corolla was supplied by a succession of overlapping green scales, from the axils of which the new buds arose. M. Germain de Saint Pierre records such a case inTrifolium repens, wherein the calyx and corolla were replaced by overlapping scales, in the axils of each one of which arose a flower; above there was a row of stamens, and in the centre a pistil in the guise of a trifoliate leaf.[139]Such instances seem to afford an extreme degree of a more common change, viz., the diminished size and contracted appearance of the sepals and petals when affected with axillary prolification. They have also a close relationship to such developments as we see in the wheat-ear carnation, in certain species of the genusMæsaand others, wherein the calyx is repeated over and again, to the partial or complete suppression of the other parts of the flower. All these cases may be in part explained by the operation of the principle of compensation.

So far as theandrœciumis concerned, the stamens either remain unaltered, or they are present in a more or less petal-like condition; but it far more frequently happens that the stamens are entirely suppressed, the adventitious bud supplying their place; thus was it in theDianthusrepresented in the adjoining woodcut, fig. 66, where the stamens were entirely absent, and their places supplied by flower-bearing branches. ThisDianthushas the more interest from its similarity to the one described by Goethe, Metam. der Pflanzen,cap. 16, sect. 105; but in that instance median prolification also existed. For my specimens I am indebted to Mr. T. Moore.

Fig.66.—Flower ofDianthussp., calyx removed; petals turned down so as to show the stalked flower-buds springing from their axils.

The pistil, too, is necessarily subject to very grave alterations when affected with this malformation. It is separated into its constituent carpels; and these assume a leaf-like aspect, and are in the great majority of instances destitute of ovules. Indeed, virescence or chloranthy is very intimately connected with this aberration, as might have been anticipated, for if the parts of the flower assume more or less of the condition of stem-leaves or bracts, it is quite natural to expect that they will partake likewise of the attributes of leaves, even at the expense of their own peculiar functions.

It occasionally happens that an adventitious bud arises from the axil of a monocarpellary pistil. This takes place sometimes inLeguminosæ, and seems to have been more frequently met with inTrifolium repensthan in other plants. The species named is, as is well known, particularly subject to a reversion of the outer whorls of the flower to leaves, and even to a leaf-like condition of the pistil. There are on record instances wherein a leaf-bud has been placed in the axil of amore or less leaf-like carpel; while at other times a second imperfect carpel has been met with in the axil of the first.[140]I have myself seen numerous imperfectly developed cases of this kind.

It may be asked whether such cases are not more properly referable to central prolification—whether the axis is not in such flowers terminated by two, rather than by one carpel? It is, however, generally admitted by morphologists that the solitary carpel ofLeguminosæis not terminal, but is the sole existing member of a whorl of carpels, all the other members of which are suppressed as a general rule, though exceptional instances of the presence of two and even of five carpels have been described.[141]

Again, the adventitious bud or carpel is placed, not laterally to the primary one, or opposite to it, on the same level, but slightly higher up—in fact, in the axil of the primary carpellary leaf. Griffith figures and describes[142]an instance of the kind in a species ofMelilotus. The stalk of the ovary is mentioned as having a sheathing base, bearing in its axil a prolongation of the axis of inflorescence, in the form of a short spike with hairy bracts and imperfect flowers, the latter having a well-formed calyx and rudimentary petals and stamens. Griffith infers, from this specimen, that the legume is not to be considered as a terminal leaf.

List of Genera in which Axillary Prolification has been observed.

OrderGenus.Leaf-bud or BranchFlower-bud or InflorescenceFrom what organ.RanunculaceæClematisFlower-budSepals.CalthaDittoDitto.AconitumDitto.DelphiniumDittoSepals, carpels, &c.Anemone!DittoInvolucre?NymphæaceæNymphæa!Fruit?NymphæaFlowerPetal.Cruciferæ*Brassica!Leaf-budFlower-budSepals and petals.Brassica!DittoStamens.Brassica!DittoDittoPistil.Cardamine!DittoSepals.Matthiola!DittoSepals and petals.Cheiranthus!DittoSepals.ErysimumDittoSepals and pistils.Lepidium!DittoPetals and stamens.ArabisDittoSepals.DiplotaxisFlower, inflorescencePistil, calyx and corolla.CapsellaCapparidaceæCleomeFlower-budSepals.Resedaceæ*ResedaDittoDitto.CaryophyllaceæArenariaBranchDitto.AgrostemmaLeaf-budDitto.*LychnisDittoStellariaDittoSileneDitto*GypsophilaDittoDittoSepals and stamens.*Dianthus!DittoDittoSepals.Dianthus!DittoInflorescencePetals and stamens.CucubalusSepalsSaponaria!Sepals and petals.MalvaceæAlceaFlower-budStamen.AurantiaceæCitrus!DittoDitto.RutaceæDictamnus!DittoPistil leafy.TropæolaceæTropæolum!DittoPetals.CelastraceæCelastrusDittoSepals.Leguminosæ*Melilotus!InflorescenceSepals and petals.MedicagoFlower-budSepals.CoronillaDittoDitto.Trifolium!DittoSecond carpel axillary to firstPistil.Melilotus!DittoDittoTrifolium!Flower-budSepals and petals.RosaceæPyrus!Fruit?Fruit?Cerasus!Flower-budPetals and stamens.Potentilla!DittoLeafy carpels.Cratægus!DittoPetals.*Rosa!DittoDittoSepals, petals, stamens and pistil.MyrtaceæLecythisDittoFruit?TetragoniaceæTetragonia?DittoDitto.CactaceæOpuntia!Fruit-like branchTufts of spines.PereskiaDittoSepals?EchinocactusDittoDitto.PhiladelphaceæPhiladelphusDittoSepals.Umbelliferæ*AthamantaDittoCalyx.*Daucus!DittoCalyx and pistil.BupleurumDittoDitto ditto.TorilisDittoDitto ditto.ApiumFlower-budCalyx and pistil.PastinacaDittoDitto ditto.Heracleum!DittoDitto ditto.Angelica!UmbelDitto ditto.Campanulaceæ*Campanula!BranchSepals.PrismatocarpusDittoFruitSepals, &c.GentianaceæGentiana!Flower-budSepals.Convolvulaceæ*Convolvulus!DittoOuter calyx.SolanaceæSolanum!DittoSepals.SolanumTubersSepals and petals.Scrophulariaceæ*Digitalis!DittoPetals, &c.VeronicaRacemeCalyx.PrimulaceæAnagallis!BranchDittoPetals.PrimulaDittoPetals and carpels.PolygonaceæRumexDittoSepals.SantalaceæThesiumLeaf-budIn place of stamens and pistils, both absent.Euphorbiaceæ?Euphorbia?Ditto?Outer bracts?OrchidaceæOrchis!Flower-budPerianth.AmaryllidaceæLeucoiumDittoDitto.IridaceæIrisDittoPistil.LiliaceæHerreriaDittoSepals.HyacinthusFlower and racemePerianth.ConvallariaFlower-budDitto.AlliumDittoDitto.CyperaceæCarexInflorescenceUtricle.

Fig.67.—Proliferous Rose. Calyx leafy; petals normal, some reflexed; stamens and pistil absent; in their places a branch with leaves and flowers.

Fig.68.—Rose exhibiting median, axillary, lateral, floral, and leafy prolification in same flower.

Complicated prolification.—From what has been before stated it may be seen that prolification of two or more kinds may coexist in the same flower. Mixed leafy and floral prolification is not unfrequent in proliferous roses, where a shoot is, as it were, prolonged through the centre of the original flower and terminated by asecond flower, or even by a cluster, as is well shown in the accompanying figure (fig. 67). Median and axillary prolification, also, not unfrequently coexist in the same flower; thus, in a proliferous rose forwarded to me by Mr. W. Thomson (fig. 68), the following changes were observed:—the swollen portion below the calyx, the "hip," was entirely absent; the sepals were leaf-like in aspect, the petals unaffected; above the petals the axis was prolonged for a short distance and then bore a circlet of miniature, sessile roses, destitute, indeed, of calyx, but provided with numerous petals, stamens, and pistils. Above these lateral flowers, the prolonged axis bore a number of scales in many rows. The scales were in their turn surmountedby a whorl of five perfect leaves, beyond which, again, the axis was prolonged into a leafy shoot terminated by a flower bud, the whole constituting a remarkably complicated admixture of elements belonging to the flower, the bud, the inflorescence, and the leafshoot.[143]

Proliferous flowers of Orchids also occasionally present great complexity in the arrangement of their parts. An instance of this kind was described by myself from specimens furnished by Dr. Moore, of Glasnevin, in the 'Journal of the Linnean Society,' vol. ix, p. 349, tabs. x, xi, and from which the following summary is extracted:

Fig.69.—Proliferous Orchis. Diagram showing the arrangement of the several organs in the seven outer circles of the flower. Each whorl is numbered, and the position of the axillary buds shown by the small circles.

The primary flowers were composed of five distinct whorls, and of at least two others less perfectly developed. These primary flowers did not give rise tomedian formations, but they produced secondary buds in the axils of the segments of the perianth. These latter buds were themselves the subject of tertiary prolification of both kinds, median and axillary. The tertiary median growths, like the primary flower, did not develop median buds, but only lateral ones—quaternary axillary prolification.

The accompanying diagrams are intended to show the plan of arrangement in these flowers. Fig. 69 shows the disposition of parts in the primary flower and the situation of the axillary buds. Fig. 70 shows the primary flower without any central prolongation, but giving off axillary buds, two of which are shown in the diagram, 2, 2; these are, each of them, the subject of both median, 3, 3, and axillary prolification, 4', 4'.

Fig.70.—Diagram to explain the construction of the double-floweredOrchis.1. The primary flower, with no median bud, the position of which, had it been present, is shown by the dotted line.2. Two axillary buds proceeding from 1, and themselves giving origin to3, 3. Median buds, and 3', 3', axillary buds.4' 4'. Axillary buds, proceeding from 3. No median bud is produced from 3; its situation, had it been present, is indicated by the dotted line.

InNarcissus majora similar combination of both forms of prolification exists, as described by Morren.[144]

On the general subject of Prolification in flowers, inaddition to the authorities already cited, the reader may refer to the following among many others:

Linnæus, 'Prolepsis,' §§ vi et vii. Goethe, 'Versuch. Metamorph.,' cap. xv and xvi §§ 103–106. Moquin-Tandon,' El. Ter. Veg.,' p. 362, &c. Engelmann, 'De Antholys.,' §§ 52–62, &c. Cramer, 'Bildungsabweichungen,' &c.Orchidaceæ,Umbelliferæ,Compositæ,Leguminosæ,Primulaceæ,Ranunculaceæ. Fleischer, 'Missbild. Cultur Gewachs.' Schlechtendal, 'Linnæa,' xv, p. 408,Rosa. 'Bot. Zeit.' vol. xx, 1862, p. 382,Cyclamen. 'Bot. Zeit.,' vol. xx, p. 301,Asphodelus; etLilium. Seringe, 'Bull. Bot.,' i, t. xi, f. 7, 8,Arabis,Diplotaxis. Clos, 'Mem. Acad. Toulouse,' 5th sér., 1862,Papaver. Wigand, 'Flora,' 1856, p. 716,Hypochæris; et 'Bot. Untersuch.,' p. 19. Buchenau,' Flora,' 1857, p. 295,Reseda. Roeper, 'Bot. Zeit.,' 1852, p. 427,Orchis. Presl., 'Linnæa,' vi, p. 599, tab. ix, figs. 5–8,Sisymbrium, Vrolik., 'Flora,' 1846, p. 97, t. i et ii, id. 1844, t. i,Digitalis. See also Schlechtendal, 'Bot. Zeit.,' vol. ix, 1851, p. 579. Klinsmann, 'Linnæa,' x, p. 604, t. v,Hesperis. Fuckel, 'Flora,' 1848, p. 609.Melilotus. De Candolle, 'Organogr.,' i, 396, t. 33. Turpin, 'Atlas de Goethe,' p. 65, t. 5, figs. 12, 13. Fenzl. 'Sitzungsbericht d. k. Akad. d. Wissensch. Wien.,' heft, iii, tabs. 3, 4,Rosa. Kirschleger, 'Flora,' 1845, 613,Dianthus,Rosa. 'Institut.,' 1841, No. 413, p. 421,Tragopogon. Baron de Melicoq., 'Ann. Sc. Nat.,' 3rd ser., vol. v. 1846, p. 61,Antirrhinum. Reichenbach, 'Icon. Fl. Germ.,' tab. 100,Reseda—"monstrosa anticipatio Euphorbiacearum et Capparidearum." Duhamel, 'Phys. Arbres.,' liv. iii, cap. 3, p. 303, pl. xii, f. 306,Rosa. Caspary, 'Bull. Soc. Bot. Fr.,' vol. vi, 1859, p. 235, Rev. Bibl.,Pyrus. Eichler, 'Flora,' 1865, tab. ix,Cleome. Lindley, 'Elements of Botany,' p. 63, &c.,Rosa,Epacris,Anagallis,Pyrus. Irmish, 'Flora,' 1858, p. 38,Pyrus; and 'Bot. Zeit.,' xix, 1861, p. 342,Hyacinthus. Duchartre, 'Bull. Soc. Bot. France,' 1861, p. 451,Rosa. Weber, 'Verhandl. Nat. Hist. Verein. Rhein. Preuss., &c.' 1858 et 1860. Landrin, 'Mem. Soc. Sc. Nat. Seine et Oise,' 1866?[145]Masters, 'Trans. Linn. Soc.,' vol. xxiii, p. 359, tab. 34 and p. 481, tab. 54.

Prolification of the embryo.—This term was applied by Moquin-Tandon to a peculiar condition of the almond (Amygdalus), in which, indeed, it is not of unfrequent occurrence. In these cases one almond encloses within its cotyledons a second embryo, and this, again, in some instances, a third, the little plants being thus packed like so many boxes one within the other. The supplementary embryos are, in the ripe state at least, quite separate and detached one from another. These cases differ from the ordinary instances wherein there is an increased number of embryos in one seed in theirposition. In the latter case, as often happens in the seeds of the orange, the new products are placed by the side one of another.[146]

For other cases of prolification or the adventitious formation of buds on leaves, roots, &c., see under Heterotaxy.

FOOTNOTES:[104]Linn., 'Prolepsis,' § vii; Goethe, 'Metamorph.,' §§ 96, 103, 106.[105]"Diaphysis inflorescentiarum." Engelmann, 'De Anthol.,' § 85.[106]'Flora,' 1844, p. 565.[107]'Ann. Sc. Nat.,' ser. 3, vol. v, 1846, p. 64.[108]'Bot Zeit.,' vol. xx, p. 382.[109]'Miscel. Austriac. Bot.,' vol. i, Vindob, 1778, p. 133.[110]"Umbellati dum prolificantur, augent umbellulam, ut ex umbellula simplici altera exeat." 'Linn. Phil. Bot.,' § 124.[111]'En. Euphorb.,' p. 36.[112]Meisner. 'Mon. Gen.PolygoniProdrom.,' p. 20, tab. v, considers the bulbils of this plant to be modifications of the pedicels of the flower.[113]See A. Braun. 'Ann. Scienc. Nat.,' 4th series, 1860, vol. xiv, p. 13.[114]"Prolificatio e latere ex calyci communi proles plurimos pedunculatos emittens, fit in compositis aggregatis proprie dictis." 'Linn. Phil. Bot.,' § 124.[115]'Bot. Zeit.,' 1857, p. 873. See also 'Verhandl. Nat. Hist. Vereins. Preuss. Rheinl. u. Westphal.,' 1854, t. ix.[116]"Pannicula spicatim sparsa onusta innumera fœtura herbaceorum flosculorum racematim cohærentium," 'Lobel. Stirp. Hist.,' p. 163. This is the "Besome Plantain, or Plantain with spoky tufts," of Ray, 'Synopsis,' p. 314. Gerard's 'Herbal,' Ed. Johnson, p. 420. Parkinson, 'Theat. Bot.,' p. 494. Baxter, 'Loudon. Mag. Nat. Hist.,' vol. ix. p. 204, and vol. iii, p. 482. fig. 118.[117]'Flora.' 1856. p. 706.[118]'Flora of Berwick-on-Tweed,' vol. i. p. 38.[119]'Adansonia,' vol. iv. 1864, p. 150, tab. vii. 'Gard. Chron.,' November 19th, 1864.[120]'Ann. Sc. Nat.,' ser. 3, tom. ii, p. 290; and 'Adansonia,' iii, tab. iv; see also Bureau, in 'Bull. Soc. Bot. France,' x, p. 191.[121]Baillon, 'Adansonia,' i, 286.[122]See also figure in 'Hort. Eystett. Ic. Plant. Vern.,' fol. 15, fig. 1.Ranunculus asiaticus.[123]Duchartre, 'Ann. des sc. nat.,' 3me série, vol. ii, 1844, p. 293.[124]Reissek, 'Linnæa,' vol. xvii, 1843, p. 641, tab. xix.[125]The tube of the calyx in these specimens was traversed by ten ribs, apparently corresponding to the primary ridges of the normal fruit; these ribs were destitute of spines, and the bristly secondary ridges were entirely absent. Those portions of the carpels which were detached from the calyx had each three ribs, a central and two lateral ones, which appeared to be continuous with the ribs of the calyx below,—although in the case of the calyx there were ten, in the case of the carpels six ribs, three to each. This diversity in number is thus explained:—A circle of vascular tissue ran round the interior of the calyx-tube, at its junction with the limb, and at the point of insertion of the petals and stamens. The vascular circle seemed to be formed from the confluence of the ten ribs from below. Of the five ribs in each half of the calyx, the three central ones were joined together just at the point of confluence with the vascular circle, above which they formed but a single rib—that traversing the centre of the carpellary leaf; the two lateral ribs of each half of the calyx seemed to be continuous, above the vascular rim, with the lateral ribs of the carpel; these lateral ribs were connected on either side with the central one by short branches of communication. The disposition of the ten ribs may be thus represented:—1 1 1 1 1 13 2 3 2 3 3 2 3 2 31 1 1 1 1 1 1 1 1 1The lower line of figures represents the calycine ribs, the middle row shows how each of these ribs is divided at the vascular rim, and the uppermost row shows their distribution above the rim. From this it will be seen that six of the calycine ribs divide into three branches, one prolonged upwards as a lateral or median rib into the carpellary leaf, the other running horizontally to join with similar branches sent out from the neighbouring rib; the four intermediate calycine ribs divide into two branches only, which join the side branches of the first mentioned, but have no direct upward prolongation into the carpel. The ten ridges are placed opposite to the sepals and petals.[126]'Neue Denkschriften derallgemeineSchweizerischen Gesellschaft,' band 5. 1841. tab. 2.[127]Bell Salter, 'Gard. Chron.,' March 13th, 1847, and 'Ann. Nat. Hist.,' 1847, vol. xix, p. 471. &c.[128]'The Origin and Production of Proliferous Flowers, with the Culture at large for raising Double Flowers from Single, and Proliferous from the Double.' By J. Hill, M.D. London, 1759.[129]A. de Candolle, 'Neue Denkschriften,' op. cit., p. 9; also Unger as cited in 'Botanical Gazette,' May, 1351. p. 70.[130]Duchartre, op. cit.[131]'Ann. Sc. Nat.,' 1844, vol. i, p. 297.[132]Maout, 'Leçons Elémentaires de Botanique,' vol. ii. p. 488; Ferrari. 'Hesperides.' pls. 271, 315, 405.[133]Moquin-Tandon, loc. cit., p. 386, &c.; see also Trécul, in the 'Bull. Soc. Bot. France,' tom. i, p, 307.[134]'Bull. Soc. Bot. Fr.,' 1863, vol. x, p. 461.[135]'Ann. Sc. Nat.,' 1835, p. 65. See also Le Maout, 'Leçons Element.,' vol. ii, p. 426.[136]'Proc. Hort. Soc.,' vol. i, p. 39, fig. 2.[137]See also 'Nat. Hist. Review,' 1865, p. 377.[138]'Acad. Roy. Belg.,' April 11th. 1863.[139]'Bull. Soc. Bot. Fr.,' tom. iii, 1856, p. 479.[140]'Linnæa,' vol. xv, p. 266,c. ic.Caspary, 'Schriften d. Physik.-Oek. Gesell. zu Königsberg,' bd. ii, p. 5, tab. iii, fig. 39, &c.[141]Lindley, 'Veg. King.,' p. 545; also Clarke on the Position of Carpels, Linn. Soc.,' December, 1850. 'Proc. Linn. Soc.,' ii, p. 105.[142]'Notulæ,' vol. i, Dicot. p. 127. 'Atlas,' pl. xliii.[143]Moquin-Tandon gives the following references to cases of proliferous roses, but some I have not been able to verify. 'Journ. des Sav.,' 22 Mai 1679. Hottinger, 'Ephem. Nat. Cur.,' dec. 3 ann. 9 et 10, p. 249. Marchant, 'Mem. Acad. Scienc. Paris.' 1707, p. 488. Preussius, 'Ephem. Nat. Cur.,' cent. 7 et 8. App. p. 83. Schuster, 'Act. Acad. Nat. Cur.,' vol. vi, p. 185. Spadoni, 'Mem. Soc. Ital.,' t. v, p. 488. See also at the end of this section for numerous other references.[144]'Bull. Acad. Belg.,' t. xx, part ii, p. 271. See also Bellynck, 'Bull. Soc. Bot. Belg.,' t. vi, ex. 'Bull. Soc. Bot. France,' t. xiv, 1867, Rev. Bibl., p. 241.Orchis ustulata.[145]I have not been able to meet with this, but it is said to contain a paper on prolification, with numerous bibliographical references.[146]'El. Ter. Veg.,' p. 364, Adnot.

[104]Linn., 'Prolepsis,' § vii; Goethe, 'Metamorph.,' §§ 96, 103, 106.

[105]"Diaphysis inflorescentiarum." Engelmann, 'De Anthol.,' § 85.

[106]'Flora,' 1844, p. 565.

[107]'Ann. Sc. Nat.,' ser. 3, vol. v, 1846, p. 64.

[108]'Bot Zeit.,' vol. xx, p. 382.

[109]'Miscel. Austriac. Bot.,' vol. i, Vindob, 1778, p. 133.

[110]"Umbellati dum prolificantur, augent umbellulam, ut ex umbellula simplici altera exeat." 'Linn. Phil. Bot.,' § 124.

[111]'En. Euphorb.,' p. 36.

[112]Meisner. 'Mon. Gen.PolygoniProdrom.,' p. 20, tab. v, considers the bulbils of this plant to be modifications of the pedicels of the flower.

[113]See A. Braun. 'Ann. Scienc. Nat.,' 4th series, 1860, vol. xiv, p. 13.

[114]"Prolificatio e latere ex calyci communi proles plurimos pedunculatos emittens, fit in compositis aggregatis proprie dictis." 'Linn. Phil. Bot.,' § 124.

[115]'Bot. Zeit.,' 1857, p. 873. See also 'Verhandl. Nat. Hist. Vereins. Preuss. Rheinl. u. Westphal.,' 1854, t. ix.

[116]"Pannicula spicatim sparsa onusta innumera fœtura herbaceorum flosculorum racematim cohærentium," 'Lobel. Stirp. Hist.,' p. 163. This is the "Besome Plantain, or Plantain with spoky tufts," of Ray, 'Synopsis,' p. 314. Gerard's 'Herbal,' Ed. Johnson, p. 420. Parkinson, 'Theat. Bot.,' p. 494. Baxter, 'Loudon. Mag. Nat. Hist.,' vol. ix. p. 204, and vol. iii, p. 482. fig. 118.

[117]'Flora.' 1856. p. 706.

[118]'Flora of Berwick-on-Tweed,' vol. i. p. 38.

[119]'Adansonia,' vol. iv. 1864, p. 150, tab. vii. 'Gard. Chron.,' November 19th, 1864.

[120]'Ann. Sc. Nat.,' ser. 3, tom. ii, p. 290; and 'Adansonia,' iii, tab. iv; see also Bureau, in 'Bull. Soc. Bot. France,' x, p. 191.

[121]Baillon, 'Adansonia,' i, 286.

[122]See also figure in 'Hort. Eystett. Ic. Plant. Vern.,' fol. 15, fig. 1.Ranunculus asiaticus.

[123]Duchartre, 'Ann. des sc. nat.,' 3me série, vol. ii, 1844, p. 293.

[124]Reissek, 'Linnæa,' vol. xvii, 1843, p. 641, tab. xix.

[125]The tube of the calyx in these specimens was traversed by ten ribs, apparently corresponding to the primary ridges of the normal fruit; these ribs were destitute of spines, and the bristly secondary ridges were entirely absent. Those portions of the carpels which were detached from the calyx had each three ribs, a central and two lateral ones, which appeared to be continuous with the ribs of the calyx below,—although in the case of the calyx there were ten, in the case of the carpels six ribs, three to each. This diversity in number is thus explained:—A circle of vascular tissue ran round the interior of the calyx-tube, at its junction with the limb, and at the point of insertion of the petals and stamens. The vascular circle seemed to be formed from the confluence of the ten ribs from below. Of the five ribs in each half of the calyx, the three central ones were joined together just at the point of confluence with the vascular circle, above which they formed but a single rib—that traversing the centre of the carpellary leaf; the two lateral ribs of each half of the calyx seemed to be continuous, above the vascular rim, with the lateral ribs of the carpel; these lateral ribs were connected on either side with the central one by short branches of communication. The disposition of the ten ribs may be thus represented:—1 1 1 1 1 13 2 3 2 3 3 2 3 2 31 1 1 1 1 1 1 1 1 1The lower line of figures represents the calycine ribs, the middle row shows how each of these ribs is divided at the vascular rim, and the uppermost row shows their distribution above the rim. From this it will be seen that six of the calycine ribs divide into three branches, one prolonged upwards as a lateral or median rib into the carpellary leaf, the other running horizontally to join with similar branches sent out from the neighbouring rib; the four intermediate calycine ribs divide into two branches only, which join the side branches of the first mentioned, but have no direct upward prolongation into the carpel. The ten ridges are placed opposite to the sepals and petals.

1 1 1 1 1 13 2 3 2 3 3 2 3 2 31 1 1 1 1 1 1 1 1 1

The lower line of figures represents the calycine ribs, the middle row shows how each of these ribs is divided at the vascular rim, and the uppermost row shows their distribution above the rim. From this it will be seen that six of the calycine ribs divide into three branches, one prolonged upwards as a lateral or median rib into the carpellary leaf, the other running horizontally to join with similar branches sent out from the neighbouring rib; the four intermediate calycine ribs divide into two branches only, which join the side branches of the first mentioned, but have no direct upward prolongation into the carpel. The ten ridges are placed opposite to the sepals and petals.

[126]'Neue Denkschriften derallgemeineSchweizerischen Gesellschaft,' band 5. 1841. tab. 2.

[127]Bell Salter, 'Gard. Chron.,' March 13th, 1847, and 'Ann. Nat. Hist.,' 1847, vol. xix, p. 471. &c.

[128]'The Origin and Production of Proliferous Flowers, with the Culture at large for raising Double Flowers from Single, and Proliferous from the Double.' By J. Hill, M.D. London, 1759.

[129]A. de Candolle, 'Neue Denkschriften,' op. cit., p. 9; also Unger as cited in 'Botanical Gazette,' May, 1351. p. 70.

[130]Duchartre, op. cit.

[131]'Ann. Sc. Nat.,' 1844, vol. i, p. 297.

[132]Maout, 'Leçons Elémentaires de Botanique,' vol. ii. p. 488; Ferrari. 'Hesperides.' pls. 271, 315, 405.

[133]Moquin-Tandon, loc. cit., p. 386, &c.; see also Trécul, in the 'Bull. Soc. Bot. France,' tom. i, p, 307.

[134]'Bull. Soc. Bot. Fr.,' 1863, vol. x, p. 461.

[135]'Ann. Sc. Nat.,' 1835, p. 65. See also Le Maout, 'Leçons Element.,' vol. ii, p. 426.

[136]'Proc. Hort. Soc.,' vol. i, p. 39, fig. 2.

[137]See also 'Nat. Hist. Review,' 1865, p. 377.

[138]'Acad. Roy. Belg.,' April 11th. 1863.

[139]'Bull. Soc. Bot. Fr.,' tom. iii, 1856, p. 479.

[140]'Linnæa,' vol. xv, p. 266,c. ic.Caspary, 'Schriften d. Physik.-Oek. Gesell. zu Königsberg,' bd. ii, p. 5, tab. iii, fig. 39, &c.

[141]Lindley, 'Veg. King.,' p. 545; also Clarke on the Position of Carpels, Linn. Soc.,' December, 1850. 'Proc. Linn. Soc.,' ii, p. 105.

[142]'Notulæ,' vol. i, Dicot. p. 127. 'Atlas,' pl. xliii.

[143]Moquin-Tandon gives the following references to cases of proliferous roses, but some I have not been able to verify. 'Journ. des Sav.,' 22 Mai 1679. Hottinger, 'Ephem. Nat. Cur.,' dec. 3 ann. 9 et 10, p. 249. Marchant, 'Mem. Acad. Scienc. Paris.' 1707, p. 488. Preussius, 'Ephem. Nat. Cur.,' cent. 7 et 8. App. p. 83. Schuster, 'Act. Acad. Nat. Cur.,' vol. vi, p. 185. Spadoni, 'Mem. Soc. Ital.,' t. v, p. 488. See also at the end of this section for numerous other references.

[144]'Bull. Acad. Belg.,' t. xx, part ii, p. 271. See also Bellynck, 'Bull. Soc. Bot. Belg.,' t. vi, ex. 'Bull. Soc. Bot. France,' t. xiv, 1867, Rev. Bibl., p. 241.Orchis ustulata.

[145]I have not been able to meet with this, but it is said to contain a paper on prolification, with numerous bibliographical references.

[146]'El. Ter. Veg.,' p. 364, Adnot.

Under this category are here included a variety of deviations from the ordinary arrangement and position of parts which cannot conveniently be classed under the preceding or under other headings. The term heterotaxy is intended to apply to the production of organs in situations where, under usual circumstances, they would not be formed. It thus does not include cases of substitution, where one part is replaced by another, or more or less metamorphosed, nor cases of multiplication, nor of prolification which are characterised not only by the production of members in unwonted situations, but also in unwonted numbers. From the very nature of the anomalies, and specially from the scanty knowledge we possess concerning their mode of development, it is not possible to allocate them in all cases correctly, and moreover many of them might as well be placed in one group as in another.

Formation of adventitious roots.—This is of exceedingly common occurrence in a vast number of plants, so much so that in most cases it cannot be considered as in any way abnormal; there are, however, a few instances where the formation of these organs may beconsidered to come within the scope of teratology, or, at least, where their production is the result of injury or of some unfavorable condition to which the plant is exposed.

Thus the production of adventitious roots on the stem of the vine is considered to be due to untoward circumstances impairing the proper action of the ordinary subterranean roots. So, too, the formation of roots on the upper portions of stems that are more or less decayed below, as in old willows, is to be considered as an attempt to obtain fresh supplies through a more vigorous and healthy channel.

A similar occurrence often arises as a consequence of some injury. Virgil had this circumstance in view when he wrote

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