CHAPTER I.

DISPLACEMENT.

Real or apparent displacement of organs from their usual position is an almost necessary consequence of, or is, at least, coexistent with a large number of teratological phenomena. It is obvious that abnormal unions or disunions, suppressions, hypertrophies, &c., are very liable to bring about or to be accompanied with changes in the position, either of the parts directly affected or of adjoining organs.

In this place, then, it is merely necessary to allude to some of the more important displacements, and to refer for further details to the sections relating to those irregularities of growth on which the displacement depends.

=Displacement of bulbs.=--I owe to the kindness of Mr. James Salter a tulip bulb which had been dug up after flowering, and from the base of which were suspended several small bulbs; and I have since seen another specimen showing the same unusual arrangement. The explanation of these formations seems to be that they correspond to the bulbils ordinarily found in the axils of the scales of the parent organ, and which, in some way or another, have been displaced and thrust into the ground. Professor de Vriese figures something of the same kind in _Ixia carminosa_.[90]

Of somewhat different nature to those above described was an anomaly described by M. Gay at a meeting of the Botanical Society of France, April 8th, 1859. The plant affected was _Leucoium æstivum_, and the changes observed were apparently attributable to a simple separation of two leaves that are usually contiguous. "Suppose," says M. Gay in describing this malformation, "the first leaf of the terminal bud separated by a long internode from the other leaves, which remain closely packed; and further, suppose an evident thickening of the upper portion of the lengthened internode, and there will be not only a single bulb, bearing with the leaves of the present year all the remnants of the leaves of the two preceding years, but two bulbs placed one above another, on the same axis, separated by the length of the internode."

The formation of bulbs in the axils of the leaves, as happens occasionally in tulips, is further alluded to under the head of hypertrophy.

=Displacements affecting the inflorescence.=--These are, for the most part, dependent on hypertrophy, elongation, atrophy, spiral torsion, &c., but there are a few instances of a different nature, which may here be alluded to as not being coincident with any of the phenomena just mentioned. Sometimes these deviations from the ordinary position have the more interest as affecting characters used to distinguish genera; thus one of the distinctions between rye-grass (_Lolium_) and wheat (_Triticum_) resides in the relative position of the spikelets and the main stem; in _Triticum_ the spikelets are placed with their backs against the rachis, in _Lolium_ with one edge against it; but in a specimen of rye-grass that has come under my own observation, the arrangement was that of _Triticum_.

M. Kirschleger relates having found a specimen of _Leucanthemum pratense_, in which the ligulate female flowers were growing singly in the axils of the upper leaves of the stem.[91] The ordinary capitulum would here seem to have been replaced by a spike or a raceme. A less degree of this change wherein a few flowers may be found, as it were, detached from the ordinary capitulum may often be observed in _Compositæ_, _Dipsacaceæ_, &c. I have also met with specimens of _Lamium album_ in which some of the fascicles or clusters of flowers in place of being placed at the same level on opposite sides of the stem were placed alternately one above another.

Caspary[92] mentions a flower of _Aldrovanda vesiculosa_, which was elevated on a stalk that was adherent to the stem for a certain distance, and then separated from it. This flower, with the leaf to which it was axillary, evidently belonged to the whorl beneath, where there was a corresponding deficiency. Another flower of the same plant bore on its pedicel a small leaf, which was doubtless the bract raised above its ordinary position.

M. Fournier mentions an instance in _Pelargonium grandiflorum_, where, owing to the lengthening of the axis, the pedicels, instead of being umbellate, had become racemose; and I owe to the kindness of Dr. Sankey a somewhat similar specimen, but in a less perfect condition. Here there was but a single flower, and that rudimentary, placed at the extremity of the axis. There were several bracts beneath this flower disposed spirally in the 1/3 arrangement, all being empty, excepting the terminal one. In like manner, a head of flowers becomes sometimes converted into an umbel.

=Displacement of leaves.=--A cohesion of parts will sometimes give rise to an apparent displacement, but the true nature of the malformation can, in general, be readily made out.

Steinheil[93] found a specimen of _Salvia Verbenaca_, the leaves of which presented very curious examples of displacement arising from cohesion. Two of these leaves placed at the base of a branch were completely fused in their lower thirds, and divided into two distinct lobes at the upper part; each of these lobes seemed to be as large as the limb of an ordinary leaf. Above these was another very broad one, apparently entire, but evidently produced by a complete cohesion of two. This completely fused leaf alternated in position with the imperfectly fused one below it; the alternation is explained by supposing that the opposite leaves of each pair were directed one towards the other, and became fused, and that thus resulted the displacement. The dislocation of the organs took place in one direction for one pair of leaves, and in another direction for the other pair, hence the alternation. Thus, leaves normally opposite and decussate may, by fusion, become alternate. A similar instance occurred to the writer in _Lysimachia vulgaris_, wherein the changes arising from fusion and suppression of parts, &c., were very considerable; as far as the leaves were concerned they presented the following arrangement in succession from below upwards:--first verticillate, then opposite, then spirally alternate, lastly opposite.[94] The term "diremption" has sometimes been applied to cases where leaves are thus apparently dragged out of position.

In _Tradescantia virginica_ I have met with opposite connate leaves; the altered position, however, being due to the union of two stems.

Twisting of the stem is a frequent cause of the displacement of leaves (see spiral torsion), as also hypertrophy, whether that excess of development take place laterally or lengthwise (see elongation). Atrophy or suppression will also frequently bring about an alteration in the position of leaves; sometimes in such a manner that the place of the suppressed organ is occupied by another one. One of the most curious instances of displacement of leaves arising from suppression is that mentioned by Morren,[95] where, in _Gesnera Geroltiana_, a large leaf apparently occupied the extremity of the axis, a position which, under ordinary circumstances, no leaf could assume. The explanation given by the Belgian professor is, that the axis in this case, instead of throwing off a pair of leaves, one on each side, had from some cause or another produced only one; this one not only being much larger than ordinary, but brightly coloured, thus assuming some of the characteristics as well as the position of the inflorescence.

Alterations in the usual arrangement of leaves, however, are not always dependent on or coexistent with other teratological changes, but may simply depend on a natural elongation of the internodes, or on fission or multiplication; for instance, in some conifers, such as the Larch, (_Abies Larix_) or _Pinus pinea_, there may be found at different stages in the growth of the branches leaves in crowded fascicles or tufts; while, when growth is more rapid, the leaves may be disposed in a spiral or alternate manner.

In the yew (_Taxus_) the leaves at the ends of the shoots not unfrequently lose their usual distichous arrangement and become arranged in a close spiral manner, the elongation of the shoot being arrested. This appears to be the result of the injury effected by some insect.

So, too, the alteration from verticillate to spiral, or _vice versâ_, may take place without any other notable change.[96] This may frequently be seen in Rhododendrons.

=Displacement of the parts of the flower.=--This subject is partly touched on in the chapters on solution, adhesion, and in those on hypertrophy, elongation, prolification, &c., so that in this place it is only requisite to offer a few general remarks, and to refer to other sections for further details. Morren, in referring to displacement of the floral organs, mentions an instance in a _Fuchsia_, wherein the four petals in place of being alternate with the sepals were placed in front of them, owing to the adhesion that had taken place between the petals and the stamens. He speaks of this transposition as metaphery.[97] The same author also gives an account of the displacement of several of the organs of the flower in _Cypripedium insigne_, the displacement being consequent, apparently, on a spiral torsion proceeding from right to left, and involving the complete or partial suppression of several of the organs of the flower. The dislocation of organs in a spiral direction led Morren to apply the term "speiranthie" to similar deviations from the usual construction. Changes of this kind among _Orchidaceæ_ are by no means uncommon; the following may be cited by way of illustration. In a specimen of _Oncidium cucullatum_ furnished me by Mr. Anderson, well known for his success as a cultivator of these plants, there was, associated with a cohesion of one sepal with another, and probably dependent on the same cause, a displacement of the sepals and petals--so that all were dragged out of place. This dislocation may be better appreciated by the accompanying formula than even by the woodcut. Let the usual arrangement be thus represented:

S

P ST P

L

S S

S standing for sepal, P for petal, L for lip, ST for stamen; then the dislocated form may be represented thus:

S

P P

T S

S_S L

In a specimen of _Cypripedium_ also furnished by Mr. Anderson the appearance was as represented in the accompanying figure and diagrams, figs. 44, 45. Referring to the plan of the natural arrangement at fig. 46, it will be seen that an explanation of the peculiar appearance of the flower may be arrived at by supposing a disunion and lateral displacement of the upper segment of the outer perianth together with the complete absence of the lower one. In the second or inner whorl of the perianth the lip is merely a little oblique on one side, but the lateral petals are distorted, displaced, and adherent one to the other and to the column, while the posterior shield-like rudimentary anther is completely wanting.

In a specimen of _Lycaste Skinneri_ similar changes were observed, as shown in the plan, fig. 47. Here the posterior sepal was deficient, the two lateral ones were present, one of them with a long tubular spur, _o o_; of the two lateral petals, _i i_, one was twisted out of place, so as partially to occupy the place of the deficient sepal; the lip was represented by two three-lobed segments, _l_, one above and within the other. The column and ovary of this flower were in their normal condition.

Cohesion of two or more segments of the perianth is frequently associated with displacements of this nature: thus, in a flower of _Dendrobium nobile_, a diagram of which is given at fig. 48, the uppermost sepal was coherent with one of the lateral ones, and at the same time diminished in size, and, as it were, dragged out of position. All the other organs of the flower are also more or less displaced, forming a minor degree of the change already alluded to, and which Morren termed speiranthy. The changes will be better appreciated by comparing them with fig. 49, a diagram showing the natural arrangement of parts in this species.

Sometimes the displacement seems consequent on hypertrophy of one of the parts of the flower, the disproportionate size of one organ pushing the others out of place. This was the case in a violet, fig. 50, in which one of the sepals _s_ was greatly thickened, and the petals and stamens were displaced in consequence.

It is curious to observe in many of these cases that the transposed organ not only occupies the place of a suppressed or abortive organ, but frequently assumes its colour, and, to some extent, its function. This has been alluded to in the case of the leaf of _Gesnera_ (see p. 88) and in Orchids this replacement seems to be very common; thus, in addition to the cases before mentioned, in a flower of an Odontoglossum, for which I am indebted to Professor Oliver, the two lateral sepals were united together and occupied the position of the labellum, which was absent. A similar occurrence happens occasionally in _Lycaste Skinneri_, thus recalling the structure of _Masdevallia_, where the labellum is normally very small. The arrangement in Lycaste may thus be symbolised:

S P st P + S S ___

[Transcriber's note: The underscores represent a horizontal curly brace in the original.]

the + indicating the position of the absent labellum.

Cases of this kind are the more interesting from their relation to the fertilization of these flowers by insects; it seems as though, when the labellum, which performs so important an office in attracting and guiding insects, is deficient, its place is supplied by other means.

Displacement of the parts of the flower from elongation of the receptacle is a not infrequent teratological occurrence, resulting sometimes in the conversion of the verticillate into the spiral arrangement. Instances of this are cited under Elongation, Prolification, &c. In this place it is merely necessary to refer to a curious circumstance that is met with in some double flowers, owing to this separation of some parts of the flower and the cohesion or adhesion of others. Thus, in some double flowers of _Primula sinensis_ and in the Pea (_Pisum sativum_), I have seen a gradual passage of sepals to petals, so that the calyx and corolla formed one continuous sheet, winding spirally around the central axis of the flower, after the fashion of a spiral tube.[98]

=Displacement of the carpels= arises from one or other of the causes above alluded to, and when suppression takes place in this whorl it generally happens that the place of the suppressed organ is occupied by one of the remaining ones, which thus becomes partially dislocated.

=Displacement of the placentas and ovules= is a necessary result of many of the changes to which the carpels are subject. The disjunction or dialysis of the carpels, for instance, frequently renders axile placentation marginal. Moreover, it frequently happens, when the carpels become foliaceous and their margins are disconnected, that the ovules, in place of being placed on the suture, or rather on the margins of the altered carpel, are placed on the surface of the expanded carpel. Thus, in some double flowers of _Ranunculus Ficaria_ that came under the writer's notice the carpels were open, _i.e._ disunited at the margins, and each bore two imperfect ovules upon its inner surface a little way above the base, and midway between the edges of the carpel and the midrib, the ovules being partly enclosed within a little depression or pouch, similar to the pit on the petals. On closer examination the ovules were found to spring from the two lateral divisions of the midrib, the vascular cords of which were prolonged under the form of barred or spiral fusiform tubes into the outer coating of the ovule. In this instance, then, the ovules did not originate from the margins of the leaf, nor from a prolonged axis, but they seemed to spring, in the guise of little buds, from the inner surface of the carpellary leaf.[99]

The occurrence, also, of different forms of placentation in different flowers on the same plant is no unusual thing in malformed flowers; thus, in double flowers of _Saponaria officinalis_ I have met with sutural, parietal, and free central placentation in the same plant.[100]

Professor Babington describes in the 'Gardeners' Chronicle,' 1844, p. 557, a curious flower of _Cerastium_, in which, in addition to other changes, the five carpellary leaves "were partially turned in without touching the placenta, which bears a cluster of ovules, and is perfectly clear of all connection with those partitions" (fig. 51). See also Lindley, 'Veg. Kingdom,' p. 497.

M. Baillon[101] records flowers of _Bunias_, some with ovules on the margins of the carpels, others with a central branch bearing the ovules; hence he concludes very justly that no fair inference can be drawn from these facts as to the normal placentation of Cruciferæ.

The same excellent observer has recorded the occurrence of free central placentation in malformed flowers of _Trifolium repens_.[102]

In malformed flowers of _Digitalis_ the change from axile to parietal placentation may often be seen. Mr. Berkeley describes an instance of this nature where the placentas were strictly parietal, and therefore receded from the distinctive characters of the order, and approximated to those of _Gesneraceæ_.

The same author alludes to certain changes in the same flower where two open carpels "were soldered together laterally, as was clear by the rudiments of two styles, the placenta being produced only at the two united edges, the outer margins remaining in the normal condition. This may possibly tend to the explanation of some cases of anomalous placentation, for the only indication of the true nature of the placentation is afforded by the two rudimentary styles, in the absence of which the spongy receptacle of the seeds must have been supposed to spring from the medial nerve."

In other cases the placentas were parietal above, but axile at the base of the capsule, a striking instance of the facility with which axile placentation becomes parietal, the change being here effected by the prolongation of the axis, and the formation on it of a second whorl of carpellary leaves.

In double flowers of _Primulaceæ_ similar alterations in the placentation may often be observed. I have seen in _Primula sinensis_ sutural, parietal, axile, and free central placentation all on the same plant; nay, even in the same capsule the ovules may be attached in various ways, and transitions from one form of placentation to another are not infrequent. The late Professor E. Forbes describes[103] an instance of true foliar and true axile placentation in the same flower in _Vinca minor_.

These and many similar changes, which it is not necessary further to allude to, are not so much to be wondered at when it is borne in mind how slight an alteration suffices to produce a change in the mode of placentation, and how frequent is the production of adventitious buds or of foliar outgrowths, as may be seen in the sections relating to those subjects and to Substitutions.

It will be remembered, also, how, in certain natural orders, under ordinary circumstances, considerable diversity in placentation exists, according as the margins of the carpels are merely valvate or are infolded so as to reach the centre. Often this diversity is due merely to the changes that take place during growth; thus, the placentation of _Caryophylleæ_, _Cucurbitaceæ_, _Papaveraceæ_, and many other orders, varies according to the age of the carpel, and if any stasis or arrest of development occurs the placentation becomes altered accordingly.

It is not necessary, in this place, to enter into the question whether the placenta is, in all cases whatsoever, a dependence of the axis, as Payer, Schleiden, and others, have maintained, or whether it be foliar in some cases, axial in others. This question must be decided by the organogenists; teratologically, however, there can be no doubt that ovules may be formed from both foliar and axial organs, and, moreover, that, owing to the variability above referred to, both in what are called natural and in what are deemed abnormal conditions, it can rarely happen that any safe inferences as to the normal or typical placentation of any family of plants can be drawn from exceptional or monstrous formations.

On the subject of placentation the following authors may be consulted:

R. Brown, 'Ann. Nat. Hist.,' 1843, vol. xi, 35. Brongniart, 'Ann. Sc. Nat.,' 1834, sér. 2. i, p. 308. Alph. De Candolle, 'Neue Denkschrift der Allg. Schweizer Gesellsch.,' Band v. 1841, p. 9. Duchartre, 'Ann. Sc. Nat.,' 3rd ser., 1844, vol. ii, p. 290. Ibid., 'Elem. Bot.,' p. 574; 'Rev. Bot.,' 1846-7, p. 213. Babington, 'Gard. Chron.,' 1844, p. 557. Lindley, 'Elements,' p. 89; 'Veg. King.,' pp. 313, 497, &c. Berkeley, 'Gard. Chron.,' 1850, p. 612. Unger, 'Nov. Act. Acad. Nat. Cur.,' 1850; and in Henfrey's, 'Bot. Gazette,' 1851, p. 70. Schleiden, 'Principles,' English edit., p. 385. Payer, 'Elem. Bot.,' pp. 196, 211, 224. Baillon, 'Adansonia.' iii, p. 310. tab. iv. Cramer, 'Bildungsabweichungen,' p. 20, &c. Clos, 'Ann. Sc. Nat.,' 5th ser., iii, 313, as well as any of the general treatises on botany. Reference may also be made to the chapters on Prolification and Substitutions (in the case of the carpels and ovules), and to the authorities therein cited.

FOOTNOTES:

[90] 'Tijdschr. voor. nat. Gesch.,' viii, 1841. tab. ii, p. 178.

[91] Communication to the Internat. Bot. Congress, Paris, 1867.

[92] 'Bot. Zeit.,' 1859, p. 117, tab. v.

[93] 'Ann. Sc. Nat.,' ser. 2, vol. iv, 1835, p. 143. tab. v.

[94] See Kirschleger, 'Flora.' 1844. p. 566 (_Scabiosa_).

[95] 'Bull. Acad. Belg.,' t. xvii. part ii, p. 387.

[96] 'Clos. Mem. Acad. Toulouse,' 5th ser., t. vi. pp. 51, 70.

[97] 'Bull. Acad. Roy. Belg.,' xviii. part ii, p. 505, and vol. xvii,