As to the causes of these structural deviations but little is known; certain of them have been already alluded to. In some cases atrophy and suppression maybe regarded as permanent states of a condition usually transitory, but this is clearly not always the case. Among external causes anything bringing about an enfeebled condition might be supposed to lead to atrophy, or suppression of some parts.

Gaertner[545] attributes the arrested development and fall of flowers to some among the following causes:--1. non-application of the pollen of the same variety, and consequent imperfect fertilisation; 2. any considerable injury to the calyx, &c.; 3. destruction of the style or stigma before the fertilisation of the ovary; 4. application to the stigma of imperfect or heterogeneous pollen or indifferent pulverulent matter; 5. defective conceptive power in the ovary.

Abortion of the ovules is considered by the same authority to be due to--1. deficiency of heat; 2. excess of moisture; 3. peculiar formation of the ovary; 4. over-luxuriant development of roots or buds; 5.

peculiar conditions of cultivation; thus, cuttings and layers produce sterile and abortive seeds much more frequently than plants of the same species raised from seed; 6. abortion of the seed is often combined with luxuriant development of the walls of the fruit.

Temperature and climatal changes in general seem not to be without effect, as has been already mentioned in the case of _Arenaria tetraquetra_, which is polygamous when growing in mountain districts.

Other ill.u.s.trations of a similar character are mentioned under the head of Heterogamy (p. 196).

Pressure has been already alluded to as one of the most obvious of the inducing causes of atrophy and suppression.

In the case of _Ranunculus auricomus_ before cited, in which the petals are rarely perfect, M. de Rochebrune considers that the deficiencies in question depend, in great measure, on the amount of moisture in the localities where the plant grows. In most places the flowers and carpels are apt to become more or less abortive, while the leaves are luxuriant; while, in dry places, the foliage is small, but the flowers are more perfect. This is quite consonant with other facts relating to the development of flowers or of leaves in general.

But while external agencies undoubtedly play some part in bringing about these changes, it is almost certain that internal causes inherent to the organization of the plant are more important. Mr. Darwin[546] accounts for the existence of rudimentary organs by the operation of the general rule of inheritance, and explains their stunted condition as the effect of disuse, not so much, of course, in the particular flower as in its predecessors. This disuse may be the result of the superior efficacy of foreign pollen as contrasted with that formed in the individual flower itself. In this way many hermaphrodite flowers tend to become dioecious, as in _Caryophyllaceae_, _Orchidaceae_, _Plantaginaceae_, _Primulaceae_ and other orders.

Although many of the circ.u.mstances above mentioned apply to plants whose structure is habitually rudimentary, there is no reason why they may not, under due restrictions, be applied to plants whose organs are only occasionally defective.

For further remarks on the subject of Abortion, the reader is referred to the sections relating to suppression, etc., also to Moquin-Tandon, 'El. Terat. Veget.,' p. 120; C. Morren, "De l'atrophie en general," in 'Bull. Acad. Belg.,' t. xviii, 1851, part i, p. 275.

FOOTNOTES:

[522] 'El. Ter. Veg.,' p. 132.

[523] _Spinosae arbores cultura saepius deponunt spinas in hortis_, 'Linn.

Phil. Bot.,' -- 272.

[524] Mr. Selby, in his 'History of British Forest Trees,' p. 465, gives the following account of the formation of this peculiar growth:--"In the autumn the parent aphis deposits her eggs at the base of the embryo leaves, within the bud destined to produce the shoots of the following year. When these begin to burst and expand in spring, the leaves, at whose bases the eggs have been deposited, instead of increasing in length, enlarge at the base, and form a cell or cyst whose mouth is at first closed by a red velvety-looking substance. If opened in this state a nest of small greenish aphides is distinctly visible, and at a certain period, or when they have acquired maturity, which is towards the end of the summer, the mouth of the cell opens and the insects fly off to inflict a similar injury upon the nascent buds of the year. In some instances the leaves of only a portion of the circ.u.mference of a shoot are affected, in which case, though a slight distortion may take place, the branch is not prevented from elongating; but in others, where the whole of the leaves around the shoot are converted into nidi, elongation is prevented and distortion to a great extent takes place."

[525] See Cramer, 'Bildungsabweich.,' pp. 53, 64, for further references.

[526] 'El. Ter. Veg.,' p. 124.

[527] Schlechtendal, 'Bot. Zeit.,' 1857, vol. xv, p. 67.

[528] On the subject of this paragraph the reader may consult A. Braun, "Ueber abnorme Blattbildung," &c., in 'Verhandl.,' d. 35, 'Naturforscherversammlung;' Jaeger, 'Flora,' 1850. p. 481, tab. 4, _Digitalis_.

[529] 'Org. Veget.,' i. p. 286.

[530] 'Bull. Soc. Bot. France, vol. viii, 1861, p. 710.

[531] 'Linnaea,' 1830, vol. v, p. 492.

[532] 'Mus. Senkenb.,' ii. p. 45.

[533] 'Bull. Acad. Roy. Belg.,' 1851, t. xviii. part i, p. 275.

[534] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 147.

[535] See Darwin, 'Variation of Domest. Anim. and Plants,' ii, 165.

[536] Gay, 'Ann. Sc. Nat.,' ser. i, 1824, t. iii, p. 44.

[537] See De Candolle, 'Mem. Legum.,' tab. 3, f. 1; Wyville Thomson, 'Trans. Bot. Soc. Edinb.,' 1851, July 10th; Berkeley, 'Gardeners'

Chronicle,' June 22nd, 1867, p. 654. A similar case is described by Dr.

Robb, in Sir W. Hooker's 'Journal of Botany,' 1841, vol. iii, p. 99, with ill.u.s.trative figures. The specimens there described were produced at New Brunswick, where plum trees flower very freely, but seldom produce ripe fruit. Dr. Robb's account is as follows:--"In the summer of 1839 I had an opportunity of watching the process of destruction among the plums, and it was as follows--Before or soon after the segments of the corolla had fallen off, the ovarium had become greenish yellow, soft, and flabby. As the fruit continued to increase in magnitude, its colour grew darker and of a more ruddy yellow, and at the end of a fortnight or three weeks the size of the abortive fruit rather exceeded that of a ripe walnut. In fact, an observer might imagine himself to be walking amongst trees laden with ripe apricots, but, like the fabled fruit on the banks of the Dead Sea, these plums, though tempting to the eye, when examined, were found to be hollow, containing air, and consisting only of a distended skin, insipid, and tasteless. By-and-bye a greenish mould is developed on the surface of the blighted fruit; then the surface becomes black and shrivelled, and at the expiration of a month from the time of flowering the whole are rotten and decomposed.

The flower appears about the beginning of June, and before August there is hardly a plum to be seen. It is curious that where two flower-stalks arise from one point of the branch, one will often go on to ripen in the normal way, while the other will become abortive, as above described."

In a specimen described by Mr. Berkeley there were two distinct ovules of equal size close to the apex of the fruit, connected with the base by vessels running down the walls. It should be observed that there is a worthless variety of plum, Kirke's stoneless, or Sans Noyau, in which the kernel is not surrounded by any bony deposit.

[538] 'Bull. Soc. Bot. Fr.,' 1862, vol. ix, pp. 37 et 291.

[539] Carl Schimp, 'Fl. Friburg,' vii, p. 745; Hook, fil., 'Journ. Linn.

Soc.,' vi, p. 9.

[540] 'Linnaea,' vol. v, 1830, p. 493.

[541] Moquin-Tandon, 'El. Ter. Veg.,' p. 325.

[542] Alph. De Candolle states that the position of the abortive ovules affords a good character for discriminating between certain species of _Quercus_, 'Bibl. Univ. Genev.,' 1862, t. xv, p. 929.

[543] See Moore, 'Nature-Printed Ferns,' 8vo, for numerous ill.u.s.trations both of depauperate and exindusiate ferns. _Scolopendrium vulgare_ seems to be one of the ferns most commonly affected in this way. Moore, loc.

cit., vol. ii, pp. 135, 147, 159, 165, &c.

[544] 'Bull. Acad. Belg.,' t. xvii, p. 38, t. 1; Lobelia, p. 85.

[545] Cited in 'Henfrey's Botanical Gazette,' i, p. 179.

[546] 'Origin of Species,' p. 450.

CHAPTER II.

DEGENERATION.

While the terms atrophy and abortion apply in the main to a mere diminution of size, as contrasted with the ordinary standard, degeneration may be understood to apply to those cases in which not only is the absolute bulk diminished, but the whole form is altered and depauperated. Degeneration, thus, is the result not so much of a deficiency in growth as of a perversion of development.

Under natural, _i.e._ habitual circ.u.mstances, the formation of pappus in place of a leafy calyx may be considered as an ill.u.s.tration of degeneration. It is evident, however, that no very decided line of demarcation can be drawn between cases of perversion and of arrest of development.

=Formation of scales.=--These may be mere epidermal excrescences, or they may be the abortive rudiments of leaves. Of this latter nature are the "cataphyllary" leaves which invest the root stocks of so many perennial plants, the perulae of leaf-buds, or the paleae on the common receptacle of composite flowers. Other ill.u.s.trations of a like character are to be met with in the membranous scales that represent leaves in _Ruscus_, _Asparagus_, _Pinus_, &c. Similar productions are met with within the flower, where they may occur as the representatives of sepals, petals, stamens, or pistils, or as mere excrescences. (See Enation.) Whole families of plants, _e.g._ _Sapindaceae_, are characterised by the presence of these organs, which are often of great interest to the morphologist as indicating the true symmetry of the flower, while they have acquired fresh importance since the publication of Mr. Darwin's work on the 'Origin of Species,' wherein we are taught to regard these rudiments as, in many cases, vestiges of organs that were more completely developed in the progenitors of the present race of plants, and the exercise of whose functions, from some cause or other, having been rendered impossible, the structures become, in process of time, proportionately stunted.

Thus, in dioecious plants we frequently find traces of stamens in the female flowers, and rudiments of the pistil in the male flower, indicating, according to the Darwinian hypothesis, that the ancestors of these plants were hermaphrodite (see Heterogamy).

Mr. Darwin has also shown that, in some cases, the utmost degree of fertility is attained, not from the action of the pollen on the stigma of the same flower, but on the influence of the male element of one blossom upon the female organs of another flower on another individual plant.

Hence, in such plants there is a tendency to a separation of the s.e.xes, while, from what has been before stated, it might be expected that rudiments of the male or female organs would be found, and also as a result of the operation of the law of inheritance. On the same principles it is easy to understand the occasional presence of the perfect in place of the rudimentary organs, as in _Dianthus_.