Chapter 9

"Quin et, caudicibus sectis, mirabile dictu,Truditur e sicco radix oleagina ligno."—'Georg.' Bk. ii.

"Quin et, caudicibus sectis, mirabile dictu,Truditur e sicco radix oleagina ligno."—'Georg.' Bk. ii.

I have seen many specimens of adventitious roots produced on the olive in the way just mentioned.

In the 'Gardeners' Chronicle,' January 8th, 1853, p. 21, is described a curious formation of roots in the fissure between two divisions of a laburnum stem. In the same journal, January 1st, 1853, p. 4, Mr. Booth mentions the case of a Cornish elm, the trunk of which was divided at the top into two main divisions, and from the force of the wind or from some other cause the stem was split down for several feet below the fork. Around the edges of the fracture, layers of new bark were formed, from which numerous roots issued, some measuring an inch in diameter and descending into the cleft portion of the tree: similar instances must be familiar to all observers.

It may happen that these roots sent down into the cavity of a decaying trunk may, after a time, become completely concealed within it, by the gradual formation and extension of new wood over the orifice of the cavity formed by the death and decay of the old wood. Such is presumed to be the explanation of a specimen ofthis kind in the possession of the writer, and taken from a cavity in an apparently solid block of rosewood; externally there were no marks to indicate the existence of a central space, but when the block was sawn up for the use of the cabinet-maker, this root-like structure was found in the centre and attached to one end of the cavity.

The production of roots which ultimately serve as props to support the branches, or as buttresses to compensate for the increasing weight of branches and foliage, is also a familiar occurrence. The huge gnaurs and burrs met with occasionally on some trees often produce great quantities, not only of adventitious buds, but of roots also.

Fig.71.—Production of adventitious roots from leaf stalk of celery.

Fig.71.—Production of adventitious roots from leaf stalk of celery.

The leaves, equally with the stems, have the power of emitting roots under certain conditions, as when the leaves are in close contact with moist soil or as the result of injury. This happens in some plants more readily than in others—Bryophyllum calycinumis a well-known instance. Mr. Berkeley has described the formation of roots from the fractured leaves of celery,[147]and also in a cabbage where a snail "having gnawed a hole into the middle of a leaf at its junction with the stem, a fascicle of roots was formed, bursting through the tissue lining the cavity, and covered with abundant delicate hairs after the fashion of ordinary radicles."

Fig.72.—Germinating plant of mango, showing production of roots from one of the cotyledons (from the Kew Museum).

Fig.72.—Germinating plant of mango, showing production of roots from one of the cotyledons (from the Kew Museum).

The production of adventitious roots is not limited to the ordinary leaves of the plant, but may be manifested on the cotyledons; thus Irmisch describes cases of this kind in the cotyledons ofBunium creticumandCarum Bulbocastanum.[148]I have figured and described an analogous case in the cotyledons of the Mango (fig. 72).[149]

To this formation of adventitious roots the gardener owes the power he has of propagating plants by cuttings,i.e., small portions of the stem with a bud or buds attached, or in some cases from portions of theleaves, of the roots themselves, or even of the fruit, as in the case of the cactus (Baillon). Care also has to be exercised in grafting certain fruit trees not to allow the grafted portion to be too close to the ground, else the scion throws out roots into the soil, and the object of the cultivator is defeated.

Figs.73 and 74 show formation of roots from leaves induced by the art of the gardener.

Figs.73 and 74 show formation of roots from leaves induced by the art of the gardener.

Layering is another garden operation dependent on the formation of these organs, and advantage is also sometimes taken of this tendency of some plants to produce roots when injured to reduce the dimensions of a plant when getting too large for the house in which it is growing. By gradually inducing the production of new roots from the central or upper portions of the stem, it becomes possible, after a time, to sever the connection between the original roots and the upper portion of the trunk, and thus secure a shortened plant.

On the subject of adventitious roots, &c., reference may be made to Trécul, 'Ann. Sc. Nat.,' 1846, t. v, p. 340, et vi, p. 303. Duchartre, 'Elements de Botanique,' p. 219. Lindley, 'Theory and Practice of Horticulture.' Thomson's 'Gardener's Assistant,' pp. 374,et seq.; and any of the ordinary botanical text-books.

On the subject of adventitious roots, &c., reference may be made to Trécul, 'Ann. Sc. Nat.,' 1846, t. v, p. 340, et vi, p. 303. Duchartre, 'Elements de Botanique,' p. 219. Lindley, 'Theory and Practice of Horticulture.' Thomson's 'Gardener's Assistant,' pp. 374,et seq.; and any of the ordinary botanical text-books.

Formation of adventitious buds on roots.—One of the characteristics by which roots are distinguished from stems in a general way consists in the absence of buds; but, as is well known, they may be formed on the roots under certain circumstances, and in certain plants, e.g.,Pyrus Japonica,Anemone Japonica, &c. What are termed suckers, owe their origin to buds formed in this situation.

If roots be exposed or injured, they will frequently emit buds. The well-known experiment of Duhamel, in which a willow was placed with the branches in the soil and the roots in the air, and emitted new buds from the latter and new roots from the former, depended on this production of adventitious organs of either kind.

Gardeners often avail themselves of the power that the roots have of producing buds to propagate plants by cuttings of the roots, but in many of these cases the organ "parted" or cut is really an underground stem and not a true root.

M. Claas Mulder has figured and described a case in the turnip-radish of the unusual formation of a leafy shoot from the root, apparently after injury.[150]From the figure it appears as if the lower portion of the root had been split almost to the extremity, while the upper portion seems to have a central cavity passing through it. From the angle, formed by the split segments below, proceeds a tuft of leaves, some of which appear to have traversed the central cavity and to have emerged from the summit, mingling with the other leaves in that situation. The production of a flower-bud has even been noticed on the root of a species ofImpatiens.

Formation of shoots beneath the cotyledons.—The tigellar or axial portion of the embryo plant, as contrasted with the radicle proper, is very variously developed in different cases; sometimes it is a mere "collar" bearing the cotyledons, while at other times it is of considerable size. Generally it does not give origin to shoots or leaves other than the seed-leaves, but occasionally shoots may be seen projecting from it below the level of the cotyledons. This happens frequently in seedling plants ofAnagallis arvensis,Euphorbiapeplus, and other species,Linaria vulgaris, someUmbelliferæ, &c.[151]

Adventitious formation of leaves.—The term phyllomania has been vaguely applied both to the production of an unwonted number of leaves and to their development in unusual situations. Under the present heading the latter class of cases are alone included. The extraordinary tendency in some Begonias to develop leaves or leafy excrescences from their surfaces is elsewhere alluded to, and is, in reality, a species of hypertrophy or over-luxuriant growth.

In some flowers where the inferior ovary is supposed to be, in part at least, formed by a dilatation of the top of the flower-stalk, leaves have been met with proceeding from the surface of the ovary or fruit, as inCratægus tanacetifolia, roses, pears, gooseberries, &c. In a specimen ofNymphæa albaI have met with scale-like leaves projecting from the surface of the fruit (or torus?), and which did not appear to be metamorphosed stamens or styles (fig. 76).

Fig.75.—Leaf proceeding from hip of the Rose.

Fig.75.—Leaf proceeding from hip of the Rose.

Fig.76.—Leaves proceeding from the ovary ofNymphæa.

Fig.76.—Leaves proceeding from the ovary ofNymphæa.

For other illustrations of increased leaf-formation, see Multiplication of foliar organs.

Fig.77.—Leontodon. Scape with two leaves; the bracts of the involucre are also leafy.

Fig.77.—Leontodon. Scape with two leaves; the bracts of the involucre are also leafy.

Production of leaves on a usually leafless inflorescence.—The development of the bracts of an inflorescence to such an extent that they resemble ordinary leaves is elsewhere alluded to as of common occurrence. It happens far less frequently that leaves are developed on an inflorescence usually destitute of them, without any metamorphosis or substitution, and without any formationof adventitious buds, such as happens in prolification. Such a partial change from a floriferous to a foliiferous branch may be seen in a specimen ofSambucus nigrain the Smithian herbarium in the Linnean Society, where the ultimate branches of the cyme bear small leaves. My attention was directed to this specimen by the Rev. W. Newbould.

Jacquin figures an analogous case inSempervivum sediforme,[152]in which the branches of the inflorescence were prolonged into leafy shoots.

Sometimes from the side of a flower-stalk or scape, which usually does not bear leaves, those organs are produced. The common dandelion,Taraxacum, sometimes offers an illustration of this, and also the daisy (Bellis).[153]In a specimen of fasciated cowslip given me by Mr. Edgeworth there was a similar formation of leaves on the flattened stalk.

Production of leaves or scales in place of flower-buds.—The position of the leaf and of the flower-buds respectively is, in most plants, well defined, but occasionally it happens that the former is formed where, under ordinary circumstances, the latter organ should be. This may happen without the formation of any transitional organs between the two, and without actual increase in the number of the buds. Where there is evidently a passage from leaf-bud to flower-bud, orvice versâ, the case would be one of metamorphy. If the number of buds be augmented, or they be mixed with the flower-buds, then it would be referable to leafy prolification of the inflorescence. There remains a class of cases wherein there is a complete substitution of one structure for the other, it may be without the slightest indication of transition between the two, and without any admixture of leaf-buds among flower-buds,or any absolute increase in the number of organs, as in Prolification. Such a case is represented in fig. 78, which shows a portion of the stem of a species ofValeriana, bearing at the summit, not an inflorescence, but a tuft of leaves without the slightest indication of flowers.

Drs. Hooker and Thomson relate that in Northern India the flowers ofAnemone rivularisare very generally absent, and their place supplied by tufts or umbels of leaves.[154]In the collection of the late Mr. N. B. Ward was a specimen of lupin in which the flowers were all absent, and their place supplied by tufts of leaves.

Fig.78.—Tuft of leaves replacing the inflorescence in a species ofValeriana.

Fig.78.—Tuft of leaves replacing the inflorescence in a species ofValeriana.

A similar appearance has been noticed inCompositæ, and I owe to the kindness of Professor Oliver the communication of a specimen of a species ofBidensfromPeru, in which the capitula, instead of consisting of florets, as usual, contained tufts of linear ciliolated bracts within the involucre, without a trace of flowers. In the eleventh volume of the 'Linnæa,' 1837, p. 301, Von Cesati figures and describes an analogous case inCarduus crispus. The same author[1] records a similar instance in the umbel ofSeseli coloratum, where the place of the flowers was occupied by stalked tufts of leaves. In the 'Gardeners' Chronicle,' October 6th, 1860, p. 894, is mentioned an instance where the blossoms of the pea were entirely absent, and their place supplied by accumulations of small, ovate, green scales, thus presenting an appearance similar to that brought about by the inordinate multiplication of the sepals in the "wheat-ear carnation," and in the Sweet William, and not unlike the condition met with inBryophyllum proliferum. InDigitalis purpureaa similar anomaly is sometimes met with.

In the apple I have observed leafy shoots bearing terminal tufts of leaves where the flower should have been, so that what, under ordinary circumstances would be a corymb of flowers, is here represented by a series of tufts of leaves. In the cultivated azaleas also, leafy shoots occupying the position of the flower may occasionally be met with.

InBouchea hyderabadensisI have seen the inflorescence more than usually branched and covered with little tufts of bracts, without a trace of true flower. A similar condition seems not infrequent inGentiana Amarella, as I have not only met with the plant myself in this condition, but have been favoured with specimens by Mr. Pamplin, Mr. Darwin, and others. InPhyteuma spicatuman analogous appearance has been recorded.

Among Griffith's collections from Affghanistan is a species of willow (Salix) in which the inflorescence replaced by a much branched panicle, bearing a quantity of minute bracts, in the axils of which nestle numerous small buds. In another specimen the inflorescencepreserves its usual catkin-like shape, but the flowers are replaced by little tufts of leaves. M. Germain de Saint Pierre mentions a case wherein the flowers ofAlisma parnassifoliawere completely replaced by leaf-buds.[155]

Fig.79.—Spikelets ofWilledenovia, composed entirely of scales to the exclusion of flowers.

Fig.79.—Spikelets ofWilledenovia, composed entirely of scales to the exclusion of flowers.

Fig.80.—Rose Willow,Salix, sp.

Fig.80.—Rose Willow,Salix, sp.

Here, also, may be mentioned the curious aggregations of scales which occur in some grasses, inRestiaceæ,Juncaceæ, and other orders, in which the inflorescenceis made up of collections of scales or bracts with no trace of floral structure. Fig. 79 shows this in a species ofWilldenovia, and a very good example is figured in a bamboo,Pseudostachyum polymorphum, by General Munro.[156]

"Rose willows" (fig. 80) owe their peculiar appearance to a similar cause, the scales of the catkin being here replaced by closely crowded leaves. These aggregations of scales or leaves are not confined to the inflorescence, but may be found in other parts of the plant, and may be frequently met with in the willow, birch, oak, &c., generally as the result of insect puncture. On the other hand, the production of leaves or leaf-buds in place of flowers is, as is well known, generally the consequence of an excess of nutrition, and of the continuance rather than of the arrest of vegetative development.[157]It has even been asserted that a flower-bud may be transformed into a leaf-bud by removing the pistil at a very early stage of development, but this statement requires further confirmation.[158]

Viviparous plants.—The spikelets of certain grasses are frequently found with some of their constituent parts completely replaced by leaves, like those of the stem, while the true flowers are usually entirely absent. A shoot, in fact, is formed in place of a series of flowers. In these cases it generally happens that the outermost glumes are changed, sometimes, however, even the outer and inner paleæ are wholly unchanged, while there is no trace of squamulæ or of stamens and pistils within them, but in their place is a small shoot with miniature leaves arranged in the ordinary manner.

The grasses most commonly affected in this manner areDactylis glomerata!,Poa bulbosa!,Poa annua!,P. trivialis!,pratensis!,alpina!,angustifolia, andlaxa,Cynosurus cristatus,Festuca nemoralis,F. ovina!,Glyceria fluitans!,Gl. aquatica,Aira alpina!,cæspitosa!,Phleum phalaroides,Lolium perenne!,Alopecurus pratensis!,Agrostis alba,Holcus mollis!

Fig.81.—Portion of panicle ofAira viviparaand separate floret.

Fig.81.—Portion of panicle ofAira viviparaand separate floret.

From an examination of the structure of viviparous grasses Von Mohl was led to the conclusion that the lower palea is to be considered as a bract, and not a perianthial leaf, because the base of the palea surrounds the stem or axis of the spikelet entirely, and both its margins cohere towards its lower extremity.[159]

A similar condition occurs not infrequently inPolygonum viviparum, and inJuncaceæ,Cyperaceæ, &c.

In the genusAlliuman analogous formation of little buds or bulbils takes place in lieu of flowers; this is specially the case withA. vineale, the flowers of which are rarely seen.

Other illustrations of a similar character, where the adventitious leaf-buds are mixed in amongst the flower-buds, are cited under the head of Prolification of the Inflorescence.

Formation of buds on leaves.—The formation of little bulbs upon the surfaces or edges of leaves, forming what are called viviparous leaves, has long been familiar to botanists amongst Alliums. Professor Alexander Braun,[160]who has paid much attention to this subject, divides cases of this kind according to the position of the buds; thus, for instance, they are sometimes formed upon the upper portion of the leaf or petiole, as in many ferns, inNymphæa guineensis, someArads, &c. The same condition has been met with as a teratological occurrence in the leaves ofCardamine pratensis,Hyacinthus Pouzolzii,Drosera intermedia,[161]Arabis pumila,Chelidonium majus,Chirita sinensis,[162]Episcia bicolor,[163]Zamia, &c.[164]Many species ofBegoniapossess the power of emitting buds from the petioles and veins of the leaf; the little ramenta or scales which so plentifully beset the surface of some of these plants likewise, in some instances, pass gradually into leaves.B. phyllomaniaca, Mart., is the species best known as manifesting this tendency, but others have it also.[165]

Buds are also very often formed upon the margins of the leaf, the best known instance of which occurs inBryophyllum calycinum; Weinmann[166]figures an instance of this kind inAlchemilla minima, or they may occur upon the lower surface of the leaf, as inOrnithogalum scilloidesandlonge-bracteatum. M. Duchartre[167]mentions a case in the tomato in which the leaves gave origin to small leaf-bearing branches, which, of course, must have originated from buds, just in the same way as in theDroserabefore mentioned.

Fig.82.—Formation of shoot on leaf ofEpiscia bicolor.

Fig.82.—Formation of shoot on leaf ofEpiscia bicolor.

Gardeners occasionally avail themselves of this formation of buds from leaves to propagate plants,e.g.Hoya,Gesnera,Gloxinia, &c.

Formation of buds in the pith.—This is said to be anormal condition in the curiousStangeria paradoxa,[168]and Mr. Berkeley records an instance of this in sea-kale[169](fig. 83) where the crown had been injured, and buds were seen sprouting from its centre.

Fig.83.—Adventitious buds in sea kale.

Fig.83.—Adventitious buds in sea kale.

Fig. 84.—Hyacinth bulb cut across to induce the formation of new bulbs.

Fig. 84.—Hyacinth bulb cut across to induce the formation of new bulbs.

Fig. 85.—Showing the formation of new bulbs on the cut edges of an old hyacinth bulb.

Fig. 85.—Showing the formation of new bulbs on the cut edges of an old hyacinth bulb.

It will be remarked that the adventitious productionof buds, like that of roots, is very often consequent on decay or injury. The Dutch bulb-growers have availed themselves of this latter circumstance in the propagation of hyacinths. Mr. Fortune, who published some articles on this subject in the 'Gardener's Chronicle,'[170]describes two special modes as adopted by these skilful horticulturists—the one to make two or three deep cuts at the base of the bulb, destroying the nascent flower-stalk when, after a time, small bulbs are formed along the edges of the cut surfaces (figs. 84, 85). The other method is effected by scooping out the interior of the base of the bulb, thus leaving exposed the cut ends of the sheathing leaves arranged concentrically; along these lines the new bulbs are, after some time, formed in great numbers (fig. 86).

Fig.86.—Showing the production of small bulbs on the inner surface of the scooped-out bulb of hyacinth.

Fig.86.—Showing the production of small bulbs on the inner surface of the scooped-out bulb of hyacinth.

For the formation of supernumerary leaves on the surface of the normal one, see Multiplication and Hypertrophy.

Production of gemmæ in place of spores.—An instance ofthis is recorded by Dr. Montagne[171]in the case of a moss,Encamptodon perichætialis, in which, in the interior of the capsule, in lieu of spores numerous minute gemmæ of the same nature as those in the cup ofMarchantiawere seen.

Formation of flowers on leaves.—It is very doubtful whether a flower-bud has ever been found actually on a leaf. Mere adhesion of the pedicels of the leaf, such as happens inRuscus, inHelwingia,Erythrochiton hypophyllanthus, and a few other plants, is, of course, not really to be considered in the light of an actual growth from the leaf, and it is very doubtful in the present state of our knowledge whether the case of the Nepaul barley should find a place here, but for convenience sake it is placed in this section, as it is uncertain at present where it properly belongs.

Fig.87.—Three-lobed end of outer palea of Nepaul barley bearing supplementary florets.

Fig.87.—Three-lobed end of outer palea of Nepaul barley bearing supplementary florets.

Fig.88.—Three spikelets of Nepaul barley.

Fig.88.—Three spikelets of Nepaul barley.

Fig.89.—Lip of outer palea of Nepaul barley.

Fig.89.—Lip of outer palea of Nepaul barley.

Fig.90.—Supplementary rachillus or outer palea of Nepaul barley bearing florets.

Fig.90.—Supplementary rachillus or outer palea of Nepaul barley bearing florets.

Fig.91.—Diagram showing arrangement of supplementary rachillus and florets.

Fig.91.—Diagram showing arrangement of supplementary rachillus and florets.

Fig.92.—Supplementary floret of Nepaul barley; palea removed.

Fig.92.—Supplementary floret of Nepaul barley; palea removed.

This curious plant has been described and figuredby Irmisch in the 13th volume of the 'Linnæa,' p. 124, t. iv; also by Professor Henslow, 'Hooker's Journal of Botany,' 1849, vol. i, p. 33, tabs. 2, 3. The lower palea of this plant forms an inverted flower-bud upon its midrib. In some fresh specimens which I have lately examined I find the structure to be as follows:—On each notch of the rachis there are three spikelets (fig. 88), each one-flowered, and each provided with two linear glumes; the outer palea in all cases is three-lobed at the summit, the central lobe being oblong and hollow, forming a kind of hood (figs. 87–89), and covered with hairs, which are directed downwards towards the centre of the plant. The two lateral lobes are morepointed than the central one; like it they are provided with hairs, but the hairs, in this case, are turned away from the centre of the plant. The cavity of the side lobes is generally empty, but that of the central lobe is occupied by a very slender stalk, which is apparently the termination of the midrib, but which is bent inwards at an acute angle, so as to occupy the hollow space (figs. 90–91). On this slender axis are developed two florets, more or less imperfect in their structure. Only one of the florets that I have seen contained a perfect ovary. The tips of the lateral lobes of the paleæ in the primary flower are sometimes extended into a long awn. A similar awn may also be occasionally found on the tips of the paleæ of the rudimentary florets. The occurrence of an adventitious axial structure with rudimentary flowers has been adduced in support of the opinion that the lower paleæ is, at least so far as its midrib is concerned, an axial rather than a foliar structure, but in the present uncertain state of our knowledge as to the morphology of grasses it is hazardous to risk any explanation founded on so exceptional a case as that of the Nepaul barley.[172]

Production of flower-buds in place of leaf-buds.—Under natural circumstances this does not appear to be of so common occurrence as the change above alluded to, but by the art of the gardener the change is often effected. In rhododendrons and in peach trees and roses I have met with this change occurring without human agency. The means adopted by the gardener are such as check the luxuriance of the leaf-shoots,[173]and this is effected in various ways, as by continuous "pinching" or removal of the leaf-buds, by pruning, ringing the bark, confining the roots, limiting thesupply of nutriment, and other means all based on the same principle. Some of the Cape bulbs (Cyrtanthus) are known not to produce their flowers till their leaves have received, in some manner, a check. Fires which often destroy the herbage thus have the effect of throwing the plant into bloom. A very remarkable instance is recorded of the production of flower-buds after an injury to the leaf-buds in the 'Bulletin of the Botanical Society of France,' vol. ix, p. 146. It appears that during the war of the French against the Arabs in Algiers, the latter planted several hundreds of Agaves with a view to obstruct the passage of the French cavalry. The soldiers hacked these plants with their sabres, and cut out the central tuft of leaves, or the heart, as gardeners call it. The following season almost every one of these Agaves sent up their large handsome flower-spikes. It is well known that, under ordinary circumstances, these plants do not flower except at long intervals of time.

Presence of flowers on spines.—That the spine, as a contracted branch, should occasionally produce flowers is not to be wondered at, though the occurrence is by no means common. M. Baillon showed at a meeting of the Botanical Society of France ('Bulletin,' vol. v, 1858, p. 316) a branched spine ofGleditschiabearing a flower at the end of each of the sub divisions. This was, therefore, strictly analogous with those cases in which the peduncle is normally spiney.

Formation of flower-bud on the petals.—An instance of this, it is believed, the only one on record, is cited in the 'Gardeners' Chronicle' for 1865, p. 760, by the Rev. M. J. Berkeley, who describes the formation of a flower-bud on the surface of a petal ofClarkia elegans. Reasoning from analogy there seems no reason why buds should not be formed on the petals as well as on the leaves.

Formation of buds on fruits.—This is a point of some moment with reference to the share which the axis takes in the production of "inferior" fruits. A very frequent malformation in pears is one wherein a second pear proceeds from the centre of the first, and even a third from the centre of the second.[174]Pears are occasionally also observed arising either from the axils of the sepals of the primary pear or from the axil of leaves originating on the outer surface of the fruits—using the term fruit in its popular sense. These cases afford strong confirmation of the view that the outer portion of the so-called fruit in these plants is rather to be considered as an expansion and hollowing-out of the flower-stalk, than as formed from the calyx-tube. It is noteworthy that the true carpels and seeds are frequently entirely absent in these cases.[175]Further reference to these fruits will be made under the head of Hypertrophy.

M. Trécul has described and figured an instance in a species ofPrismatocarpus, in which a second flower proceeded from the axil of a bract attached to the side of the fruit of the first flower.[176]A similar growth was observed in the fruit ofPhiladelphus speciosusby M. A. Gris, who observed that the so-called calyx-tube was provided with two small bracts, from the axil of one of which proceeded a small flower-bud.[177]

Fig.93.—Small buds projecting from the edges of the fruit inOpuntia.

Fig.93.—Small buds projecting from the edges of the fruit inOpuntia.

The fruits ofOpuntia Salmiana,O. fragilis,[178]O. monacantha, and of some species ofEchinocactus, have been observed to form small fruit-like branches around their summits. M. Napoléon Doumet describes the fruit as ripening as usual, but as being destitute of seeds in the interior; after a little while the fruit begins to wither,and then a circle of small buds, like those of the stem, may be seen at the top of the fruit, each bud springing from the axil of a little tuft of wool and spines found on the fruit. These little buds elongate into long shoots, produce flowers the following year, which flowers exhibit the same peculiarity. Gasparini and Tenore are said to have recorded the same fact as long since as 1832. The specimen from which the figure (fig. 93) was taken produced its fruits in the Royal Gardens at Kew, and is now preserved in the museum of that establishment. The adventitious growth in these cases appears to arise from the tufts of spines, which, it has been suggested, are the homologues of the sepals. There can, however, be little doubt that the outer and lower portion of the fruit ofOpuntiaand its allies is a dilatation of the flower-stalk. This is borne out by the fruits ofPereskia, which bear leaves on their surface arranged spirally; indeed, the fruits ofPereskia Bleoare mentioned as producing buds from theirsummits, in the same way as theOpuntiajust cited.P. Bleois said, by M. Delavaud,[179]to present this anomaly as a constant occurrence. On the summit of the primary fruit, arising apparently from the axils of the sepals, or of small leafy bracts in that situation, are a series of fruit-like branches, which, in their turn, are surmounted by others, even to the fourth generation.

The fruits ofTetragonia expansafrequently have attached to their side a secondary flower or fruit in such a position as to lead to the inference that it springs from the upper portion of the peduncle which is dilated to invest the true carpels. In other instances it is due to an adhesion of the pedicel to the side of the fruit. In either case the production of an adventitious bud might be considered as an illustration of prolification of the inflorescence, though not as was supposed by Moquin and others of axillary prolification.[180]

Buds have also been produced artificially on the surface of some of the fruits in the construction of which the axis is supposed to share; thus, the unripe fruits of some species ofLecythiswere stated by Von Martius, at a meeting of the German Naturalists at Carlsruhe, to produce buds when placed in the earth. The fruit of these plants is probably of the same nature as that of thePomaceæ, and Baillon[181]succeeded in producing buds on the surface of the inferior ovary ofJussiæa.

Some of the cases just mentioned have been considered to be instances of prolification of the fruit, but the fruit has little to do with the appearances in question.

Formation of adventitious flowers and fruits within the ovary.—This generally arises either from substitution of aflower-bud for an ovule or from prolification; there are certain cases, however, where the new growth seems not to be either due to metamorphosis or to prolification strictly.

The cut, fig. 94, represents a case where, in the dilated upper portion of the ovary ofSinapis arvensis, two flower-buds were found projecting from a raised central line, corresponding, as it would seem, to the midrib, and not to the margins of the carpel. Similar cases have occurred inNasturtium amphibium,Brassica Rapa, andPassiflora quadrangularis.

Fig.94.—Distended pod ofSinapis arvensisbearing in the interior stalked flower buds.

Fig.94.—Distended pod ofSinapis arvensisbearing in the interior stalked flower buds.

In Bromfield's 'Flora Vectensis,' p. 35, the following account is given of an abnormal development inCardamine pratensis: "On the lower part of the corymb were several seed vessels on pedicels changed from their usual linear to an ovate elliptical figure, so as to resemble a silicula. These, on being opened, were found to contain petals of the usual colour, which in the pods above had burst from their confinement and appeared as semi-double flowers; the valves of the pod answering to the true calyx. * * * From their verticillate arrangement it is evident that these petaloid expansions were not transformed seeds, but simply a development of the common axis within the ovary into an abortive whorl of floral organs, besides which there were evident rudiments both of stamens and germensin the centre of the bundle." Baillon[182]also records a case of the same nature inSinapis arvensis.

Fig.94*.—Portion of the interior of the silicle inCheiranthus Cheiri, showing adventitious pod in the place of an ovule.

Fig.94*.—Portion of the interior of the silicle inCheiranthus Cheiri, showing adventitious pod in the place of an ovule.

Fig.95.—Adventitious pod from fig. 94, enlarged.

Fig.95.—Adventitious pod from fig. 94, enlarged.

Here, too, may also be mentioned the presence of an adventitious siliqua within the ordinary one attached along the same line as the ovules, and partially divided by a replum into two cavities. In this case there was nothing to indicate the presence of floral envelopes (figs. 94, 95). A similar occurrence has been brought under my notice in some grapes which were observed to be cracking before they were perfectly ripe, and in which adventitious fruits were found within the parent grape, occupying the position of seeds (figs. 96, 97).

Similar anomalous growths are noticed under the heads of Substitution and Prolification.

Formation of stamens within the cavity of the ovary.—The only instance of this that has come under the author's observation occurred in some flowers ofBæckea diosmæfolia,Rudge, for an examination of which he is indebted to Mr. Bentham.

Fig.96.—Section of Barbarossa grape showing adventitious grape in the position of a seed.

Fig.96.—Section of Barbarossa grape showing adventitious grape in the position of a seed.

Fig.97.—Grape with supplementary fruit in the interior

Fig.97.—Grape with supplementary fruit in the interior

In the normal flower there is a turbinate hollow calyx, whose limb is divided into five serrated lobes; alternating with these latter, and springing from the throat of the calyx, are the petals. Originating from the same annular disk as the petals are the stamens, seven or eight in number. The ovary is partially adherent, is surmounted by a style, and has two or three loculi with an axile placenta, to which several small curved ovules are attached. The malformed flowers did not present anything peculiar in their outer parts, nor did the ovary, partially immersed within the expanded top of the flower-stalk and the calyx-tube, which is continuous with that organ, show externally any indication of the change within. On cutting it across, however, in any direction, numerous perfect stamens (filaments and anthers) were seen projecting from the walls of the cavity (fig. 98). In most of the flowers the ovary was one-celled; but in a few there was the usual axile placenta; yet even in these latter cases the stamens originated from the walls of the cavity, and not from the placenta. The stamens presented different degrees of development; in some cases they were fully formed, the anther-lobes open, and the pollen exposed; while in other instances thefilaments were involute or circinate, just as the ordinary stamens are in the unexpanded flower-bud. In some cases imperfect stamens were found, mere barren filaments, with or without rudimentary anthers at the top. In no instance was there a perfect ovule, or, indeed, any trace of ovules. The stamens appeared to be arranged irregularly on the walls of the ovarian cavity; and while they were certainly more numerous at the lower portion (that now generally considered to be formed by the cup-like end of the pedicel), they were not wanting in the upper half of the ovary (or that which is probably formed from the carpellary leaves).


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