_Crataegus Oxyacantha_, _Dianthus sinensis_, _Daucus Carota_, _Cerasus
Lauro-cerasus_. De Candolle alludes to a case of the kind in the bean,
and figures a species of _Solanum_ with three cotyledons.[420] Jaeger
alludes to a similar instance in _Apium Petroselinum_;[421] Ehrenberg to
one in the marigold (_Calendula_);[422] Reinsch to an analogous
appearance in the beech (_Fagus_), associated with a union of the
margins of two out of the three cotyledons, and of those of two out of
the three leaves next adjacent.[423] This fusion seems frequently to
accompany increase in the number of cotyledons. It was so in the
_Correa_, and in the _Crataegus_ previously mentioned. Some of these
cases may be accounted for by chorisis or by a cleavage of the original
cotyledons, as happens, according to Duchartre,[424] in some Coniferae,
which he considers to be improperly termed polycotyledonous. Whether
this holds good in the Loranths, where (_Nuytsia_, _Psittacanthus_) an
appearance of polycotyledony exists, is not stated. In the case of the
rue (_Ruta_) figured by M. A. de Jussieu[425] this splitting of one
cotyledon into two is sufficiently evident, as is also the case in the
sycamore (_Acer pseudo-platanus_), seedlings of which may often be met
with divided cotyledons.
In other instances a fusion of two embryo plants may give rise to a
similar appearance, as in the _Euphorbia_ and _Sinapis_ found by M.
Alph. de Candolle (see _ante_, p. 56).
=Pleiotaxy or multiplication of whorls.=--In the preceding section
notice has been taken of the increased number of parts in a single
whorl, but an augmentation of the number of distinct whorls is still
more frequently met with. Many of the so-called double flowers owe their
peculiarity to this condition. The distinction between the two modes in
which the parts of the flower are increased in number has been pointed
out by Engelmann, Moquin, and others, and the two seem to require
distinctive epithets; hence the application of the terms polyphylly and
pleiotaxy, as here proposed.
=Pleiotaxy in the bracts.=--An increase in the number of bracts has been
met with very constantly in a species of _Maesa_, and in a peculiar
variety of carnation, called the wheat-ear carnation.[426] In some of
these cases the increase in the number of bracts is attended by a
corresponding suppression in the other parts of the flower. Such a
condition has been frequently met with in _Gentiana Amarella_, where the
bracts are increased in number, coloured purple, and destitute of any
true floral organs. A similar condition exists in some varieties of
_Plantago major_ (var. _paniculata_), as has been previously stated, p.
109.
[Illustration: FIG. 187.--Wheat-ear carnation. The appearance is due to
the multiplication of the bracts and the suppression of the other parts
of the flower.]
It has been noticed also in the common pea, _Pisum sativum_, and M.
Lortet[427] records a case of the kind in _Erica multiflora_, the
flowers of which, under ordinary circumstances, are arranged in
clusters, but in this case the pedicels were more closely crowded than
usual, and were covered for their whole length with small rose-coloured
bracts arranged in irregular whorls, the upper ones sometimes enclosing
imperfect flowers. In the 'Gardeners' Chronicle,' 1865, p. 769, is
figured a corresponding instance of _Delphinium Consolida_, in which the
bracts were greatly increased in number, petaloid, and, at the same
time, the central organs of the flower were wholly wanting.
[Illustration: FIG. 188.--_Delphinium Consolida_. Multiplication of
bracts at the expense of the other parts of the flower.]
[Illustration: FIG. 189.--Multiplication of bracts, &c., _Pelargonium_.]
In flowers of _Pelargonium_ may occasionally be seen a repetition of the
whorls of bracts, in conjunction with suppression and diminished size of
some of the other portions of the flower (fig. 189).
The common foxglove (_Digitalis purpurea_) has likewise occasionally
been observed subject to a similar malformation.
_Cornus mas_ and _C. suecica_ sometimes show a triple involucre.[428]
Irmish[429] records an analogous case in _Anemone Hepatica_, wherein the
involucre was doubled. Similar augmentation occurs in cultivated
Anemone. In addition to the plants already mentioned, Engelmann[430]
mentions as having produced bracts in unwonted numbers, _Lythrum
Salicaria_, _Plantago major_, _Veronica spicata_, _Echium vulgare_,
_Melilotus arvensis_, and _Rubus fruticosus_.
It must here be remarked that this great number of the bracts occurs
naturally in such plants as _Godoya_, in which the bracts, or, as some
consider them, the segments of the calyx, are very numerous, and
arranged in several overlapping segments.
In some of the cultivated double varieties of _Nigella_ the finely
divided involucral bracts are repeated over and over again, but on a
diminished scale, to the exclusion of all the other parts of the flower.
=Pleiotaxy or repetition of the calyx.=--The true calyx is very seldom
affected in this manner, unless such organs as the epicalyx of mallows,
_Potentilla_, &c., be considered as really parts of the calyx.
In _Linaria vulgaris_ Roeper observed a calyx consisting of a double
series, each of five sepals, in conjunction with other changes.[431] It
is also common in double columbines, delphiniums, nigellas, &c.
In the 'Revue Horticole,' 1867, p. 71, fig. 9, is described and figured
by M. B. Verlot a curious variety of vine grown for years in the Botanic
Garden at Grenoble, under the name of the double-flowered vine. The
place of the flower is occupied by a large number of successive whorls
of sepals disposed in regular order, and without any trace of the other
portions of the flower. It is, in fact, more like a leaf-bud than a
flower. The outermost whorls of this flower open at the time when the
ordinary flowers of vines do; the second series are gradually produced,
and expand about the time when the ovaries of the normal flowers begin
to swell; a third series then gradually forms, and so on, until frost
puts a stop to the growth. This malformation, it appears, is produced
annually in certain varieties of vine, and may be perpetuated by
cuttings.
The flower of the St. Valery apple, already alluded to under the head of
sepalody, might equally well be placed here. It is not very material
whether the second whorl of organs be regarded as a repetition of the
calyx or as a row of petals in the guise of sepals.
Engelmann[432] cites the following plants as occasionally presenting a
repetition of the calyx, in most cases with a suppression of the other
floral whorls:--_Stachys lanata_, _Myosotis palustris_, _Veronica
media_, _Aquilegia vulgaris_, _Nigella damascena_, _Campanula
rapunculoides_.
=Pleiotaxy in the perianth.=--Increase in the number of whorls in the
perianth is common in lilies, narcissus, hyacinths, &c. It may be also
met with occasionally among orchids. The lily of the valley
(_Convallaria maialis_) seems also to be particularly subject to an
increase in the number of parts of which its perianth consists, the
augmentation being due partly to repetition or pleiotaxy, partly to the
substitution of petaloid segments for stamens and pistils.[433]
In this place may also be mentioned the curious deviation from the
ordinary structure occasionally met with in _Lilium candidum_, and known
in English gardens as the double white lily. In this case there are no
true flowers, but a large number of petal-like segments disposed in an
irregular spiral manner at the extremity of the stem, some of the
uppermost being occasionally verticillate.[434]
[Illustration: FIG. 190.--Double white lily. Multiplication of
perianth-segments and other changes.]
=Pleiotaxy of the corolla.=--With reference to double flowers, it was
remarked by Linne that polypetalous flowers were, as he said,
multiplied, while monopetalous flowers were duplicated, or triplicated,
as the case may be,[435] a statement that is true in the main, though
it requires modification. In the case of polypetalous, or rather
dialypetalous flowers, the petals may be very largely increased by
multiplication, as in roses, anemones, pinks, &c. In the last-named
genus the number is often so much increased that the calyx splits from
the tension exercised on it by the increasing mass within. This
multiplication may happen without any metamorphy or substitution of
petals for stamens, though, in the majority of cases, it is associated
with such a change. It is curious to observe in some of these flowers
that the total number of parts is not greatly increased; thus, in some
of the double-flowered _Leguminosae_, such as _Ulex europaeus_ and _Lotus
corniculatus_, the petals are repeated once or twice, the stamens are
petalodic, but reduced in number, while the carpels are usually entirely
wanting. Thus, owing to the diminished number of parts in the inner
whorls of the flower, these very double-looking blooms do not contain
any greatly increased number of parts.[436]
Flowers that, under ordinary circumstances, are gamopetalous, become, in
some instances, multiplied by the formation of additional segments, just
as in the case of polypetalous corollas; but in these cases the corollas
become polypetalous, their petals do not cohere one with another. Among
double flowers of this character may be mentioned _Campanula
rotundifolia_, _Gardenia_ sp., _Nerium Oleander_, _Serissa_ sp.,
_Arbutus Unedo_, &c. The change is associated with petalody of the
stamens and pistils.
A more frequent change among the monopetalous orders is the duplication
or triplication of the corolla, in consequence of which there appear to
be a series of corollas enclosed one within the other, the lobes of
which generally alternate with one another, but which sometimes are
superposed. This happens occasionally in the primrose (_Primula
acaulis_), and constitutes the variety called by the gardeners "hose in
hose."
The same condition occurs frequently in some species of _Datura_ and
_Campanula_.
[Illustration: FIG. 191.--_Campanula rotundifolia_. "Double flowers"
resulting from dialysis and multiplication of the petals.]
In _Antirrhinum majus_ double flowers of this character sometimes occur;
the outermost corolla is normal, the succeeding ones usually have their
petals separate one from the other; the stamens are sometimes present,
sometimes absent, and at other times petalodic. Similar occurrences may
be met with in labiates and jasmines, and in _Erica hyemalis_.
Mr. W. B. Hemsley has kindly furnished me with flowers of a similar kind
occurring in wild specimens of _Epacris impressa_,[437] and there are
analogous phenomena in the common honeysuckle (_Lonicera Periclymenum_),
in which three corollas and no stamens often occur.
This duplication may either be accounted for on the theory of chorisis
above alluded to, or by supposing that the extra corolline whorl is due
to a series of confluent petalodic stamens; that the latter is the true
explanation, in certain cases at least, is shown by some flowers of
_Datura fastuosa_, in which the second corolla was partially staminal in
its appearance, and bore nearly perfect anthers, in addition to the five
ordinary stamens, which were unaltered either in form or position. Some
partially virescent honeysuckle flowers have a similar structure.
There are other cases of apparent multiplication or duplication, due,
probably, rather to the formation of outgrowths from the petals than to
actual augmentation of their number. These excrescences occur sometimes
on the inner surface of the petals, or of the corolla; at other times on
the outer surface, as in some gloxinias, &c. This matter will be more
fully treated of under the head of hypertrophy and enation.
=Pleiotaxy of the androecium.=--An increase in the number of whorls in
the stamens is very common, especially in cases where the number of
circles of stamens is naturally large. The augmentation of the number of
stamens is still more frequent where these organs are arranged, not in
verticils, but in one continuous spiral line.
In _Cruciferae_ there is always an indication of two whorls of stamens,
and this indication is rendered even more apparent in some varieties
accidentally met with. So in _Saponaria_, in _Dianthus_, and other
_Caryophylleae_, three and four verticils of stamens have been met with.
In _Lonicera Periclymenum_ a second whorl of stamens more or less
petalodic sometimes occurs.
Moquin mentions a variety of _Rubus fruticosus_ in which nearly 900
petaloid organs existed in the place of the twenty-five or thirty
stamens natural to the plant, the other organs of the flower being in
their ordinary condition, with the exception of the pistil, which did
not attain its full size. Baillon records the occasional existence of
two rows of stamens in _Ditaxis lancifolia_.
=Increased number of stamens in orchids, &c.=--Various deviations from
the ordinary type of orchid structure have been already alluded to under
the head of displacement, fusion, peloria, substitution, &c., but the
alterations presented by the androecium in this family are so
important in reference to what is considered its natural conformation,
that it seems desirable, in this place, to enter upon the teratological
appearances presented by the androecium in this order, in somewhat
greater detail than usual. The ordinary structure of the flower with its
three sepals, two petals, labellum, column; and inferior ovary, is well
known. Such a conformation would be wholly anomalous and inexplicable
were it not that the real number and arrangement of parts have been
revealed by various workers labouring to the same end in different
fields. Thus, Robert Brown, Link, Bauer, Darwin, and others, paid
special attention to the minute anatomy and mode of distribution of the
vessels; Irmisch, Crueger, Payer, and others, to the evolution of the
flower; Lindley, St. Hilaire, and Reichenbach, to the comparison of the
completed structures in the various genera and species; while the
teratological observers have been numerous, as will be seen from the
selected references cited at the end of this paragraph and in other
places. The result of this manifold study has been a pretty general
agreement that the structure of the order (omitting minor details) is as
follows:--A six-parted perianth in two rows, the outer three (sepals)
generally regular and equal in shape; of the inner three (petals or
tepals) two are regular, and one, the labellum very irregular,
consisting not only of a petal, but of two abortive stamens incorporated
with it. The column is considered to be made up of one perfect and three
abortive stamens, in inseparable connection with three styles. By some,
however, it is supposed that all the stamens are confluent with the
column and none with the lip.
[Illustration: FIG. 192.--Diagram showing the arrangement of parts in an
orchid flower. According to Crueger, the stamens A 2, A 3, should be
distinct from the lip. The uppermost figure 2 should have been 1. (See
text.)]
In either case it is admitted that there are six stamens in two rows.
The first row consists of one posterior stamen, which is generally
perfect, and two abortive stamens incorporated with the labellum. The
second row also consists of three stamens, all of which are usually
abortive and inseparable from the column. Traces of them may
occasionally be met with in the form of tubercles or wing-like processes
from the column. In _Cypripedium_, while the ordinary stamen of the
outer row is deficient, two of the inner series are present. The
diagram, fig. 192, will serve to show the arrangement of the parts as
above described. + represents the situation of the stem or axis; on the
opposite side is the bract; between these are placed the sepals, one
posterior or next the axis (incorrectly numbered 2 in the plan), two
lateral 1, 1; next in order follow the petals, 2, 2, 2, two lateral and
somewhat posterior, one larger (the lip), anterior; the outer series of
stamens are represented by A 1, A 2, A 3, the two latter being fused
with the labellum; _a_ 1, _a_ 2, _a_ 3 represent the position of the
inner verticil of stamens, while s, s, s denote the three carpels. It is
foreign to the purpose of this book to detail the varied evidence in
support of this explanation of the homologies of orchid flowers.[438]
All that can be done in these pages is to set forth the evidence
furnished by teratology as to this matter--evidence for the most part
accumulated and recorded without any special reference to any theory of
orchid structure.
The following details all refer to flowers in which the number of
stamens in orchidaceous plants was increased beyond what is necessary.
They are arranged with reference to the number of adventitious organs,
beginning with those in which the number was smallest, and proceeding
thence to those in which it was greatest. In some cases it has not been
possible to ascertain whether the adventitious organs were really
restorations of the numerical symmetry, substitutions of one part for
another, stamen for petal, &c., or wholly adventitious productions.
Unless otherwise stated, the interpretation put upon the facts thus
recorded is that of the present writer, and not necessarily that of the
original observer.
Mr. J. T. Moggridge has described and figured a flower of
_Ophrys insectifera_ in which there was a vestige of a second
stamen present, probably one of the inner series fig. 192
(_a_^2).[439] The same observer also records the presence of a
second anther between the lobes of the normal one. This can
hardly be referred to either of the typical stamens, but would
seem to be a perverted development of the rostellum.[440]
Roeper is stated by Cramer[441] to have seen a specimen of
_Orchis morio_ with two stamens.
In a flower of _Habenaria chlorantha_, described by the late
Professor Henslow,[442] the outer three stamens are suppressed,
while two of the inner group are present, as happens normally
in _Cypripedium_.
A flower of _Cattleya violacea_ afforded a similar
illustration; but in this case only one of the inner stamens
was developed, and this in the form of a small petal, partly
adherent to the column.
In _Dendrobium normale_, Falconer, not only is the perianth
regular, but the column is triandrous,[443] the three stamens
(according to the diagram of its structure given by Lindley)
pertaining to the outer row.
In a specimen of _Dendrobium hoemoglossum_ kindly forwarded
from Ceylon by Mr. Thwaites there were three stamens present,
of which one posterior belonged to the outer series A 1, and
two lateral to the inner _a_ 1, _a_ 2, fig. 192.
M. His observed, several years in succession, some flowers of a
species of _Ophrys_ with three sepals, no lateral petals, one
lip, and three perfect stamens. In this case probably the two
supernumerary stamens were petals which had assumed an
anther-like character.
Wydler describes a flower of _Ophrys aranifera_ in which one
outer and two inner stamens were present.[444] I have myself
met with three such flowers in the same species. The stamens
present were A 1, _a_ 1, _a_ 2.
Dr. J. E. Gray exhibited at the Botanical Society of London, in
August, 1843, a specimen of _Ophrys apifera_ with a triandrous
column, the supernumerary anthers belonging, apparently, to the
inner whorl.
In his 'Catalogue of the Plants of South Kent,' p. 56, tab. iv,
f. 16, the Rev. G. E. Smith describes and figures a flower of
_O. aranifera_ with a triandrous column, seemingly of the same
kind as that spoken of by Dr. Gray.
Mr. Moggridge met with a triandrous flower in the same species,
and refers the appearance to "a fusion of two flowers,
accompanied by suppression and modification."[445] As, however,
no details are given in support of this opinion, it may be
conjectured that the two additional stamens were members of the
inner whorl _a_ 1, _a_ 2, and thus the conformation would be
the same as in the flowers just mentioned. The figures given by
Mr. Moggridge bear out this latter view, while they lend no
support to the hypothesis advanced by him. Nevertheless, no
decided opinion can be pronounced by those who have not had the
opportunity of examining the flowers in question.
Alphonse de Candolle[446] figures a flower of _Maxillaria_ in
exactly the same condition, so far as the stamens are
concerned, as in the Ophrys flowers just mentioned. It is
curious to observe that in many of these cases the two lateral
petals are suppressed.
Von Martius mentions the occurrence of three anthers
(_naturaliter conformatae_) in _Orchis morio_.[447] Richard,
as cited by Moquin-Tandon, Lindley, and others, describes and
figures a peloria of _Orchis latifolia_ with regular triandrous
flowers.[448]
The writer has examined, in the Royal Gardens at Kew, a flower
of _Cattleya crispa_ in which were three stamens, the central
one normal; the two lateral ones, belonging probably to the
inner whorl, were in appearance like the lateral petals, and
one of them was adherent to the central perfect column.
Duchartre[449] mentions a flower of _Cattleya Forbesii_ in
which there were two labella in addition to the ordinary one,
the column being in its normal condition. From the analogy of
other cases it would appear as if the additional labella in
this instance were the representatives of two stamens of the
outer whorl. Beer likewise has put on record the existence of a
triandrous _Cattleya_.[450]
A specimen of _Catasetum eburneum_ forwarded by Mr. Wilson
Saunders was normal so far as the sepals and two lateral petals
were concerned, but the anterior petal or labellum was flat and
in form quite like the two lateral ones; the column was normal
and in the situation of the two anterior stamens of the outer
series A 2, A 3, were two labella of the usual form (fig. 156,
p. 291). Perhaps the _Oncidium_ represented at p. 68, fig. 29,
may also be explained on the supposition that the two lateral
lobes of the labellum in this flower were the representatives
of stamens.
In Fig. 193 is shown the arrangement of parts in a flower of
_Ophrys aranifera_. Here there were three sepals, two lateral
petals, one of which was adherent to the side of the column;
the central labellum was seemingly deficient, but there were
two pseudo-labella placed laterally in the position of the two
antero-lateral stamens of the outer series (A 2, A 3). Within
these was another perfect stamen occupying the position of the
anterior stamen of the inner series (_a_ 3). In another flower
of the same species, gathered at the same time (fig. 194),
there were three sepals not at all different from those of the
normal flower. The three petals next in succession were also,
in form and position, in their ordinary state. In colour,
however, the two upper lateral petals differed from what is
customary, in having the same purplish-brown tint which
characterises the lip. Within these petals, at the upper part
of the flower, there was the ordinary column, and at the
opposite side, alternating with the petals before mentioned,
two additional lip-like petals, one provided with a half-anther
containing a single perfectly formed pollen-mass (A 2, A 3). It
is, perhaps, worthy of notice that the arrangement of the
coloured spots on the true labellum, and that on the
adventitious lips, replacing the two lower of the outer
stamens, were not of a similar character. The supernumerary
lips had the [Greek: pi]-shaped marking which is so common in
this species, while the true lip was, as to its spots, much
more like _O. apifera_. Alternating with this last whorl were
three columns, all apparently perfectly formed and differing
only from the ordinary one in their smaller size and
corresponding to _a_ 1, _a_ 2, _a_ 3. The ovary in this flower
was two-celled, with four parietal placentas, thus giving an
appearance as though there had been a fusion of two or more
flowers associated with suppression and other changes. The
position of the supernumerary organs and the absence of any
positive sign of fusion in the bracts or other part of the
flower, seemed, however, to negative the idea of fusion.[451]
[Illustration: FIG. 193.--Diagram showing the arrangement of
parts in a malformed flower of _Ophrys aranifera_ (see p.
384).]
[Illustration: FIG. 194.--Malformed flower of _Ophrys
aranifera_ with two supernumerary lips and three additional
stamens.]
A similar illustration, for a knowledge of which the writer is
indebted to the kindness of Professor Asa Gray and Mr. Darwin,
occurred in some specimens of _Pogonia ophioglossoides_
collected by Dr. J. H. Paine in a bog near Utica, New York. It
will be seen from the following description that these flowers
presented an almost precisely similar condition to those of the
_Ophrys aranifera_ just mentioned. "The peculiarities of these
flowers," writes Professor Gray, "are that they have three
labella, and that the column is resolved into small petaloid
organs. The blossom is normal as to the proper perianth, except
that the labellum is unusually papillose, bearded almost to the
base. The points of interest are, first, that the two accessory
labella are just in the position of the two suppressed stamens
of the outer series, viz. of A2 and A3, as represented in the
diagram, fig. 192; and there is a small petaloid body on the
other side of the flower, answering to the other stamen, A1.
Secondly, in one of the blossoms, and less distinctly in
another, two lateral stamens of the inner series (_a_1 and
_a_2) are represented each by a slender naked filament. There
are remaining petaloid bodies enough to answer for the third
stamen of the inner series and for the stigmas, but their order
is not well to be made out in the dried specimens." It may here
be mentioned that _Isochilus_ is normally triandrous.
A tetrandrous flower of _Cypripedium_ has also been recorded.
In _Isochilus_, according to Cruger, there are often five
stamens, and there are several, besides those already
mentioned, in which six more or less perfect stamens have been
seen--of these the following may be taken as illustrations. A
hexandrous flower of _Orchis militaris_ has been recorded by
Kirschleger,[452] and in the accompanying diagram (fig. 195),
from Cramer,[453] of a monstrous flower of _Orchis mascula_,
there is one perfect stamen of the outer row and two lip-like
stamens of the same series, while the inner verticil comprises
one perfect and two abortive stamens.
[Illustration: FIG. 195.--Diagram of flower of _Orchis mascula_
with two additional lips, two perfect and two imperfect stamens
(after Cramer).]
Morren[454] describes some flowers of _Orchis morio_ in which
there were three sepals, three petals, and within the latter
two other ternary series of petals; this would seem to be a
case of petalody of all six stamens. Morren, however, seems to
have considered the additional segments as repetitions of the
corolline whorl, though he describes a central mass as the
column bearing a "_souvenir_ of the anther." Nevertheless,
there is no decisive evidence either in his figure or his
description in support of his opinion as to the nature of the
central mass, which might be a distorted condition of the
styles, or, as is more probable, a rudimentary and irregular
flower. Morren also describes another flower of the same plant
in which there were three sepals, two lateral petals partially
lip-like in aspect, a third labellum normal, two additional
labella representing the two anterior stamens of the outer
whorl, while more or less developed rudiments of the remaining
four stamens also exist.
While, in most cases, the supernumerary stamens can, by reason
of their relative position, their complete or partial
antheriferous nature, be safely referred to one or other of the
six stamens, making up a typical orchid flower, there are other
specimens in which the additional stamens are altogether
adventitious, and do not admit of reference to the homologue.
Thus it was in a specimen of _Odontoglossum Alexandrae_ examined
by the writer, and in which, within a normally constructed
perianth, there were six columns, all polliniferous, but
arranged in so confused and complicated a manner that it was
impossible to make out any definite relation in their position.
There was nothing to indicate a fusion of flowers, but rather
an extension of the centre of the flower, and consequent
displacement of the stamens, &c. Again, the existence of
adventitious stamens does not necessarily imply the development
of organs usually suppressed, inasmuch as they may result from
the assumption by the lateral petals of staminal
characteristics.
Nevertheless, as far as teratology is concerned, specimens may
be found in which some or all of the usually suppressed stamens
of _Orchidaceae_ may be found. These stamens may be all perfect
(polliniferous), or, as is more frequently the case, more or
less petal-like. Moreover, when the stamens are petalodic, the
form assumed is usually that of the labellum.
The presence of stamens in undue numbers in orchids is very
generally, but not always, attended by some coincident
malformation, of which the most frequent is cohesion of two or
more sepals, and consequent displacement or adhesion of one
petal to the side of the column. Petalody of the styles and
median prolification are also sometimes found in association
with an augmented number of stamens.
[Illustration: FIG. 196.--Increased number of carpels, tulip.]
[Illustration: FIG. 197.--Fruit of St. Valery apple cut lengthwise.]
=Pleiotaxy of the gynoecium.=--An increase in the number of whorls of
which the pistil consists is not of very frequent occurrence. Generally
after the formation of the whorl of carpels, the energy of the growing
point ceases, or if by chance it be continued, the result is more
generally the production of a new flower-bud (median prolification) than
the repetition of the carpellary series. It is necessary also to
distinguish between the veritable augmentation of the pistil and the
semblance of it, brought about by the substitution of carpels for some
other organs, as pistillody of the stamens, and even of the segments of
the perianth, is not very unfrequent, as has already been stated under
the head of substitution. Again, the increased number of carpels which
is sometimes met with in such flowers, as _Magnolia_ or _Delphinium_,
where the ovaries are arranged in spiral series, is not strictly
referable to the present category.
The orange is one of the plants most frequently subject to an
augmentation in the number of carpellary whorls; sometimes this is due
to the stamens assuming the guise of carpels, but at other times the
increase occurs without any alteration in the stamens or other organs.
If the adventitious carpels be exposed, they are covered with yellow
rind, while those portions that are covered by the primary carpels are
destitute of rind. Some varieties of the double tulip are very subject
to a similar change, but, in this case, the petals and the stamens very
frequently become more or less carpellary in their nature. Fig. 196
represents an increased number of whorls of carpels in the variety
called "rex rubrorum," the segments of the perianth having been removed.
In the St. Valery apple, already referred to, there is a second whorl of
carpels above the first, a fact which has been made use of to explain
the similar structure of the pomegranate.
The tomato (_Lycopersicum esculentum_) is another plant in which an
adventitious series is frequently produced, and generally in combination
with the primary series.
In the Chinese primrose (_Primula sinensis_) a supernumerary whorl is
frequently met with, generally associated with other changes in the
construction and arrangement of the parts of the flower.
M. de Candolle[455] mentions a flower of _Gentiana purpurea_ with four
carpels in one series, and five others in the circle immediately above
them. Wigand[456] alludes to an instance wherein there was a second pair
of carpels above the first in _Vinca herbacea_. Dr. Sankey has forwarded
flowers of a _Pelargonium_ having a double series of carpels, eight in
the outer row, five in the inner, and this condition is stated to exist
in the flowers of the same plant for two years consecutively. In
_Aquilegia_ I have met with a similar increase in the whorls of
carpels.[457] Meissner records a similar augmentation in _Polygonum
orientale_.[458]
Wigand[459] describes and figures a flower of _Vinca minor_, in which
there were two carpels intervening between the ordinary pair, and a
similar illustration has been observed by the writer in _Allamanda
cathartica_. Eichler[460] has put on record a similar case in a
capparid.
Marchand[461] mentions a polycarpellary berberid (_Epimedium
Musschianum_). The supernumerary carpels in this flower were placed on a
short axis, which originated in the axils of the stamens, and as these
latter organs were present in their usual number and position, the
adventitious carpels could not be considered as resulting from a
transformation, or substitution of carpels for stamens.
Lastly, the instance cited by Dr. Allman[462] in _Saxifraga Geum_ may be
alluded to. Here there was a row of adventitious carpels between the
stamens and pistils, the backs of the carpels being turned towards the
axis of the flowers. Dr. Allman explains the presence of the
supernumerary parts by the supposed production of a whorl of secondary
axes between the stamens and the centre of the flower. These axes are
further supposed to bear imperfect flowers, of which the additional
carpels are the only traces, but this explanation seems forced.
In addition to the references already cited the following may be given:
Duchartre, 'Ann. Sc. Nat.,' 4 ser., vii, p. 23 (Tulip).
Ferrari, 'Hesperides,' pp. 271, 395, 405. Duchartre, 'Ann. Sc.
Nat.,' 4 ser., 1844, vol. i, p. 294. Maout, 'Lecons Element.,'
vol. ii, pp. 488-9. Clos, 'Ann. Sc. Nat.,' 1865, p. 317
(_Citrus Aurantium_).
Clos, 'Bull. Soc. Bot. Fr.,' vol. xiii; 'Rev. Bibl.,' p. 75.
Pasquale, 'Reddicont Accad. Sc. Fis. e Math. Napoli.' Octr.
1866 (_Solanum Lycopersicum_).
On the general subject of multiplication, in addition to previous
citations, the reader may be referred to A. P. de Candolle, 'Theorie
Element. Bot.,' ed. 3, p. 89.
=Increased number of flowers in an inflorescence.=--This happens
generally as a result of over luxuriant growth, and scarcely demands
notice here, being rather referable to variation than to malformation.
The increased number of florets in the spikelets of some grasses has
already been alluded to (p. 351). Thus spikelets of wheat occasionally
produce more than the three florets which are proper to them.[463] It
will be remembered that in this as in many other grasses there are
rudimentary florets, and it is no matter for surprise that these florets
should occasionally be fully developed.
FOOTNOTES:
[392] 'Bull. Soc. Bot. Fr.,' vol. vii, 1860, p. 587.
[393] 'Fragment. Phyt. Austral.,' part xx, p. 270.
[394] 'Bull. Acad. Belg.,' xvi, pt. i, p. 60, "Fuchsia," p. 125, c. ic.
[395] "Theorie de la feuille," 'Arch. des Sciences Bibl. Univers.,'
1868.
[396] See Engelmann, 'De Antholysi,' p. 16, section 12.
[397] Verhandl. des Botanisch. Vereins Brandenburg,' 1859, 1 heft.
[398] See Henslow. 'Mag. Nat, Hist.' 1832, vol. v, p. 429.
[399] 'Phytologist,' September, 1857.
[400] Seemann's 'Journal of Botany,' iv, p. 168, t. 47, f. 3.
[401] 'Illust. Hortic.,' 1866, misc., p. 97.
[402] See Fresenius, 'Mus. Senkenb.,' bd. 2, p. 43. Schlechtendal, 'Bot.
Zeit.,' iv, pp. 403, 492, _Veronica tetrandra_.
[403] 'Flora,' 1865, tab. 6, fig. 8.
[404] 'Org. Veget.,' t. i, p. 497, pl. 42, f. 3.
[405] 'El. Ter. Veg.,' p. 354.
[406] Cited in "Rev. Bibl." of 'Bull. Soc. Bot. Fr.,' 1866, p. 171.
[407] Loc. cit., 351.
[408] 'Mem. Acad. Toulous.,' vi, 1862, ex 'Bull. Soc. Bot. Fr.,' "Rev.
Bibl.," vol. ix, 1862. p. 127.
[409] 'Flora.' 1857. p. 289.
[410] L. c., p. 354.
[411] Giraud, 'Ed. Phil. Mag.,' Dec., 1839.
[412] See _Cerasus Caproniana_, D. C. 'Plant. Rar. Hort. Genev.,' tab.
18.
[413] Nees, 'Linnaea,' v, p. 679, tab. 11 (_Schoenodorus_).
[414] 'Gard. Chron.,' 1852, p. 452.
[415] See Cramer, 'Bildungsabweich.' pp. 16, 24.
[416] 'Linnaea,' 1842, p. 389, c. ic.
[417] Seemann's 'Journal of Botany,' 1867, vol. v, p. 158.
[418] Cramer, 'Bildungsabweich,' p. 66, _Astrantia major_, _Eryngium_,
to which may be added _Daucus_, _Heracleum_, &c.
[419] See also Reinwardt, 'Nov. Act. Acad. Nat. Cur.,' 12, 1, 37; and
Masters, 'Journ. Linn. Soc.,' vi, p. 24.
[420] 'Organog. Veget.,' tab. 53.
[421] 'Missbild.,' p. 206.
[422] Ehrenberg, 'Flora,' 1846, p. 704.
[423] 'Flora,' 1860, tab. 7.
[424] 'Ann. Sc. Nat.,' 3 ser., t. x, p. 207.
[425] 'Mem. Mus.,' xii. t. 17.
[426] 'Nov. Act. Acad. Nat. Cur.,' xv, tab. xxviii, f. 3; 'Bot. Mag.,'
t. 1622. "Caryophyllus spicam frumenti referens." A similar malformation
in _Dianthus barbatus_ is not uncommon. It has lately been introduced
into gardens under the name of _Dianthus_ "_mousseux_," but is not
likely to find favour with gardeners.
[427] 'Bull. Soc. Bot. France,' t. vi, 1859. p. 268.
[428] Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' 1860.
[429] 'Bot. Zeit.,' 1848, p. 217.
[430] 'De Anthol.,' p. 17, Sec. 12.
[431] 'Linnaea,' vol. ii, 1827, p. 85.
[432] 'De Antholysi,' p. 17, tab. iii, f. 15, 16; Weinmann, 'Phytanth.
iconogr.,' nro. 292.
[433] See Hildebrand, 'Bot. Zeit.,' 1862, p. 209, tab. viii; Cramer,
'Bildungsabweich.,' p. 7, tab. xiii; Engelmann, 'De Antholysi,' p. 18,
&c. For similar changes in _Gagea arvensis_ see Wirtgen, 'Flora,' 1838,
t. xxi. p. 350, and 'Flora.' 1846, p. 353. Some of these are cases of
synanthy.
[434] Schlechtendal, 'Bot. Zeit.,' xx, 1862, p. 301.
[435] 'Phil. Bot.,' Sec. 126.
[436] C. Morren, 'Bull. Acad. Belg.,' xix, part ii, p. 17.
[437] 'Seemann's Journal of Botany,' iii, p. 354.
[438] On this point the reader will find an excellent summary in
Lindley's 'Vegetable Kingdom,' cd. iii, p. 183_a_, and in Darwin,
'Fertilisation of Orchids,' p. 292. See also Crueger,'Journ. Linn. Soc.,'
t. viii, p. 134.
[439] 'Seemann's Journal of Botany,' vol. iv, p. 168, tab. 47.
[440] Ibid., t. iv. 1866, p. 168, t. xlvii, f. 1.
[441] 'Bildungsabweich,' p. 8; see also 'Bot. Zeit.,' 1852, p. 425.
[442] 'Journ. Linn. Soc.,' t. ii, p. 104. tab. 1, fig. B.
[443] Lindl., "Orchid. Ind.," 'Jour. Linn. Soc.,' iii, p. 9.
[444] 'Arch. Bot.,' ii, p. 300, tab. xvi, f. 11.
[445] 'Seemann's Journal of Botany,' v, p. 318, tab. lxxii, figs. A 4, 4
_a_.
[446] "Monstr. Veg.," in 'Neue Denkschrift,' p. 17, tab. vii.
[447] 'Flora,' t. viii, 1825, p. 736.
[448] 'Mem. Soc. d'Hist. Nat.,' ii, 1, p. 212, tab. iii.
[449] 'Bull. Soc. Bot. Fr.,' t. vii, 1860, p. 26.
[450] 'Beitr. Morphol. und Biol. Orchid.,' quoted by Cramer;
'Bildungsabweich,' p. 9.
[451] Masters, 'Journ. Linn. Soc.,' viii, p. 207. See also Rodigas,
'Bull. Soc. Bot. Belg.,' iv, p. 266, for similar changes in _Cypripedium
Hookerae_.
[452] Kirschleger, 'Flora,' 1844, p. 131.
[453] 'Bildungsabweich,' p. 11, tab. xiv, f. 3.
[454] 'Bull. Acad. Roy. Belg.,' t. xix, part 2, p. 171.
[455] 'Organogr. Veget.,' t. i, p. 509, tab. 40, figs. 6, 7.
[456] 'Flora,' 1856, p. 715.
[457] 'Linn. Trans.,' t. xxiii, p. 364, tab. 34, fig. 5.
[458] 'Monog. Polygon,' pl. 3, K. f. 12.
[459] 'Flora,' 1856, tab. viii.
[460] Ibid., 1865, tab. ix, f. 6.
[461] 'Adansonia,' vol. iv, 1864, p. 127.
[462] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 126.
[463] See Schlechtendal, 'Bot. Zeit.,' t. xviii, p. 381 (_Triticum_);
also 'Flora,' t. xiv, 1831, p. 5 (_Avena_).
PART II.
DIMINISHED NUMBER OF ORGANS.
A diminution in the number of parts is generally due to suppression,
using that word as the equivalent of non-development. It corresponds
thus in meaning with the _Fehlschlagen_ of the Germans, the _avortement
complete_ of Moquin and other French writers. It differs from atrophy,
or partial abortion, inasmuch as the latter terms apply to instances
wherein there has been a partial development, and in which evolution has
gone on to a certain extent, but has, from some cause or other, been
checked. These cases will be found under the head of diminished size of
organs. As the word abortion is used by different authors in different
ways, it is the more necessary to be as precise as possible in the
application of the term. In the present work abortion is used to apply
to cases wherein parts have been formed, but wherein growth has been
arrested at a certain stage, and which, therefore, have either remained
_in statu quo_, while the surrounding parts have increased, or have,
from pressure or other causes, actually diminished in size.
In practice, however, it is not always possible to discriminate between
those instances in which there has been a true suppression, an absolute
non-development of any particular organ, and those in which it has been
formed, and has grown for a time, but has afterwards ceased to do so,
and has been gradually obliterated by the pressure exercised by the
constantly increasing bulk of adjacent parts, or possibly has become
incorporated with them. In the adult flower the appearances are the
same, though the causes may have been different.
CHAPTER I.
SUPPRESSION OF AXILE ORGANS.
Absolute suppression of the main axis is tantamount to the non-existence
of the plant, so that the terms "acaulescent," "acaulosia," etc, must be
considered relatively only, and must be taken to signify an atrophied or
diminished size of the stem, arising from the non-development of the
internodes.
The absence of lateral branches or divisions of the axis is of frequent
occurrence, and is dependent on such causes as the following:--deficient
supply of nutriment, position against a wall or other obstacle, close
crowding of individual plants, too great or too little light, too rich
or too poor a soil, &c.
Probably the absence of the swollen portion below the flower in the case
of many proliferous roses, double-flowered apples, as already referred
to, may be dependent on the non-development of the extremity of the
peduncle or flower-stalk. Thus, in a double-flowered apple recently
examined, there was a sort of involucel of five perfect leaves, then
five sepals surrounding an equal number of petals, numerous stamens, and
five styles, but not a trace of an expanded axis, nor of any portion of
the carpels, except the styles. The views taken as to the nature of this
and similar malformations must depend on the opinion held as to the
nature of inferior pistils, and on the share, if any, that the expanded
axis takes in their production. As elsewhere said, the evidence
furnished by teratology is conflicting, but there seems little or
nothing to invalidate the notion that the end of the flower-stalk and
the base of the calyx may, to a varying extent, in different cases,
jointly be concerned in the formation of the so-called calyx-tube and of
the inferior ovary. Obviously it is not proper to apply to all cases
where there is an inferior ovary the same explanation as to how it is
brought about.
As these pages are passing through the press, M. Casimir de Candolle has
published a different explanation as to the nature of the hip of the
rose, having been led to his opinion by the conclusion that he has
arrived at, that the leaf is to be considered in the light of a
flattened branch, whose upper or posterior surface is more or less
completely atrophied.
According to M. de Candolle, the calyx-tube, in the case of the rose, is
neither a whorl of leaves, nor a concave axis in the ordinary sense in
which those terms are used, but is rather to be considered as a
ring-like projection from an axis arrested in its ulterior development.
The secondary projections from the original one correspond to an equal
number of vascular bundles, and develope into the sepals, petals,
stamens, and ovaries. If these organs remained in a rudimentary
condition, the tube of the calyx would be reduced to the condition of a
sheathing leaf. The rose flower, then, according to M. de Candolle, may
be considered as a sheathing leaf, whose fibro-vascular system is
complete, and from which all possible primary projections are
developed.[464]
If, as M. de Candolle considers, the leaf and the branch differ merely
in the fact that the vascular system is complete in the latter, and
partly atrophied in the former, it would surely be better to consider
the "calyx-tube" of the rose as a concave axis rather than as a leaf,
seeing that he admits the fibro-vascular system to be complete in the
case of the rose.
With reference to this point the reader is referred to Mr. Bentham's
account of the morphology and homologies of the _Myrtaceae_ in the
'Journal of the Linnean Society,' vol. x, p. 105. See also _ante_, pp.
71, 77.
Some doubts also exist as to the nature of the beak or columella of such
fruits as those of _Geraniaceae_, _Malvaceae_, _Umbelliferae_,
_Euphorbiaceae_, &c. The nature of the organ in question may probably be
different in the several orders named; at any rate the subject cannot be
discussed in this place, and it is mentioned here because, now and then,
it happens that the organ in question is completely wanting, and hence
affords an illustration of suppression.
FOOTNOTES:
[464] 'Theorie de la feuille.' p. 24.
CHAPTER II.
SUPPRESSION OF FOLIAR ORGANS.
This subject may be considered, according as the separate leaves of the
stem or of the flower are affected, and according as either the number
of members of distinct whorls, or that of the whorls themselves, is
diminished.
The terms aphylly, meiophylly, and meiotaxy may be employed, according
as the individual leaves are altogether wanting, or with reference to
the diminished number of parts in a whorl, or a decrease in the
verticils.
=Aphylly.=--Entire suppression of the leaves is a rare phenomenon. Under
ordinary circumstances it occurs in most _Cactaceae_, in some of the
succulent Euphorbias, and other similar plants, where the epidermal
layers of the stem fulfil the functions of leaves. But even in these
plants leaf-like organs are present in some stage or another of the
plant's life.
Partial suppression of the leaf occurs sometimes in compound leaves,
some or other of the leaflets of which are occasionally suppressed.
Sometimes, as Moquin remarks, it is the terminal leaflet which is
wanting, when the appearance is that of _Cliffortia_, at other times the
lateral leaflets are deficient, as in _Citrus_ or _Phyllarthron_.
_Ononis monophylla_ and _Fragaria monophylla_ may be cited as instances
of the suppression of the lateral leaflets. If the blade of the leaf
disappears entirely, we have then an analogous condition to that of the
phyllodineous acacias.
With reference to the strawberry just mentioned, Duchesne, 'Hist. Nat.
Frais.,' p. 133, says that this was a seedling raised from the _fraisier
des bois_, and the characters of which were reproduced by seed, and have
now become fixed. The monophyllous condition has been considered to be
the result of fusion of two or more leaflets, but however true this may
be in some cases, it is not the case with this strawberry. M. Paillot
states that he has found the variety in a wild state.[465]
In like manner varieties of the following plants occur with simple
leaves, _Rosa berberifolia_ (_Lowea_), _Rubus Idaeus_, _Robinia
pseudacacia_, _Fraxinus excelsior_, _Sambucus nigra_, _Juglans nigra_,
&c.
In one instance seen by the writer every portion of the leaf of a rose
was deficient, except the stipules and a small portion of the petiole.
(See abortion.)
=Meiophylly.=--A diminished number of leaves in a whorl occasionally
takes place; thus, in some of the _Stellatae_, and frequently in _Paris
quadrifolia_, the number of leaves in the verticil is reduced. Care must
be exercised in such instances that an apparent diminution arising from
a fusion of two or more leaves be not confounded with suppression.
=Meiophylly of the calyx or perianth.=--A lessened number of sepals is
not a very common occurrence among dicotyledonous plants. Seringe
figures a proliferous flower of _Arabis alpina_ with two sepals only,
and a similar occurrence has been noticed in _Diplotaxis tenuifolia_.
In _Cattleya violacea_ the writer has met with a flower in which the
uppermost sepal was entirely wanting, while two of the lateral petals
were fused together. Moquin records that in some of the flowers of
_Chenopodiaceae_, in which the inflorescence is dense, a suppression of
two or three sepals sometimes occurs. The species mentioned are _Ambrina
ambrosioides_, _Chenopodium glaucum_, and _Blitum polymorphum_.
=Meiophylly of the corolla.=--Suppression of one or more petals is of
more frequent occurrence than the corresponding deficiency in the case
of the sepals. Among _Caryophyllaceae_ imperfection as regards the
numerical symmetry of the flower is not uncommon, as in species of
_Cerastium_, _Sagina_, _Dianthus_, &c. In _Ranunculaceae_ the petals are
likewise not unfrequently partially or wholly suppressed. A familiar
illustration of this is afforded by _Ranunculus auricomus_, in which it
is the exception to find the corolla perfect.[466] Some varieties of
_Corchorus acutangulus_ in west tropical Africa are likewise subject to
the same peculiarity. Amongst _Papilionaceae_ absence of the carina or of
the alae is not uncommon, as in _Trifolium repens_, _Faba vulgaris_, &c.
Moquin relates a case of the kind in the haricot bean, in which the
carina was entirely absent, and another in the pea, where both carina
and alae were missing, thus reducing the flower to the condition that is
normal in _Amorpha_ and _Afzelia_. Suppression of the upper lip in such
flowers as _Calceolaria_ has been termed by Morren "apilary."
In _Orchidaceae_ entire absence of the labellum, frequently without any
other perceptible change, is of common occurrence. The writer has seen
numerous specimens of the kind in _Ophrys apifera_ and _O. aranifera_;
also in _Dendrobium nobile_, _AErides odoratum_, _Cypripedium villosum_,
_Listera ovata_, &c. Morren[467] mentions analogous deficiencies in
_Zygopetalum maxillare_, _Calanthe_ sp., and _Cattleya Forbesii_. In
most of these there was also a fusion of the two lower sepals, which
were so twisted out of place as to occupy the situation usually held by
the labellum. At the same time the column was partially atrophied. To
this deficiency of the lip the author just quoted proposed to apply the
term acheilary, [Greek: a-cheilarion]. Mr. Moggridge has communicated to
the author an account of certain flowers of _Ophrys aranifera_, in which
the petals were deficient, sometimes completely, at other times one or
two only were present.
=Meiophylly of the androecium.=--Suppression of one or more stamens,
independently of like defects in other whorls, is not uncommon, even as
a normal occurrence, _e.g._ in _Carlemannia_, where the flower, though
regular, has only two stamens, and other similar deficiencies are common
in Dilleniads.
Seringe relates the occurrence of suppression of some of the stamens in
_Diplotaxis tenuifolia_,[468] St. Hilaire in _Cardamine hirsuta_, others
in _C. sylvatica_.
In _Caryophyllaceae_ suppression of one or more stamens has been observed
in _Mollugo cerviana_, _Arenaria tetraquetra_, _Cerastium_, &c.[469]
Among violets the writer has observed numerous flowers in which two or
three stamens were suppressed. Chatin[470] alludes to a similar
reduction in _Tropaeolum_, while in flowers that are usually didynamous
absence of two or more of the stamens is not unfrequent, _e.g._ in
_Antirrhinum_, _Digitalis_, while in a flower of _Catalpa_ a solitary
perfect stamen, and a complete absence of the sterile ones usually
present, have been observed. This might have been anticipated from the
frequent deficiencies in the staminal whorl in these plants under what
are considered to be normal conditions. Reduction of the staminal whorl
is also not unfrequent in _Trifolium repens_ and _T. hybridum_, and has
been seen in _Delphinium_, &c.[471]
=Meiophylly of the gynoecium.=--Numerical inequality in the case of
the pistil, as compared with the other whorls of the flower, is of such
common occurrence, under ordinary circumstances, that in some text-books
it is looked on as the normal condition, and a flower which is isomerous
in the outer whorls is by some writers not considered numerically
irregular if the number of the carpels does not coincide with that of
the other organs.
But in this place it is only necessary to allude to deviations from the
number of carpels that are ordinarily found in the particular species
under observation. As illustrations the following may be
cited:--_Arenaria tetraqueta_, which has normally three styles, and a
six-valved capsule, has been seen with two styles, and a four or
five-valved capsule. Moquin relates an instance in _Polygala vulgaris_
where there was but a single carpel, a condition analogous to that which
occurs normally in the allied genus _Mozinna_. _Reseda luteola_
occasionally occurs with two carpels only, while Aconites, Delphiniums,
Nigellas, and Paeonies frequently experience a like diminution in their
pistil.
In a flower of _Papaver Rhaeas_ the writer has recently met with an ovary
with four stigmas and four parietal placentae only, and to Mr.
Worthington Smith he is indebted for sketches of crocus blooms with two,
and in one instance only a solitary carpel.
Moquin cites the fruit of a wild bramble (_Rubus_) in which all the
little drupes which go to make up the ordinary fruit were absent, except
one, which thus resembled a small cherry. In _Crataegus_ the pistil is
similarly reduced to a single carpel, as in _C. monogyna_.
The writer has on more than one occasion met with walnuts (_Juglans_)
with a single valve and a single suture.[472] If the ovary of _Juglans_
normally consisted of two valvate carpels, the instances just alluded to
might possibly be explained by the suppression of one carpel, but the
ovary in _Juglans_ is at first one-celled according to M. Casimir de
Candolle.
Among monocotyledons _Convallaria majalis_ may be mentioned as very
liable to suffer diminution in the number of its carpels, either
separately or in association with other changes.[473]
=Meiophylly of the flower as a whole.=--In the preceding sections a
reduction in the parts of each individual whorl has been considered
without reference to similar diminution in neighbouring verticils. It
more commonly happens, nevertheless, that a defect in one series is
attended by a corresponding imperfection in adjoining ones. Thus
trimerous fuchsias and tetramerous jasmines may frequently be met with,
and Turpin describes a tetramerous flower of _Cobaea scandens_. Perhaps
monocotyledonous plants are more subject to this numerical reduction of
the parts of several verticils than are other flowering plants. Thus, in
both _Lilium lancifolium_ and _L. auratum_ the writer has frequently met
with pentamerous flowers. In _Convallaria maialis_ a like deviation not
unfrequently occurs.[474] M. Delavaud has recorded a similar occurrence
in a tulip.[475]
Dimerous crocuses may also sometimes be met with. In one flower of this
nature the segments of the perianth were arranged in decussating pairs,
and the four stamens were united by their filaments so as to form two
pairs.
M. Fournier mentions something of the same kind in the flower of an
_Iris_.[476]
Orchids seem peculiarly liable to the decrease in the number of their
floral organs. Prillieux[477] mentions a flower of _Cattleya
amethystina_ wherein each whorl of the perianth consisted of two
opposite segments.
The same observer has put on record instances of a similar kind in
_Epidendrum Stamfordianum_. In one flower of the last-named species the
perianth consisted of one sepal only, and one lip-like petal placed
opposite to it.[478] Morren[479] describes a flower of _Cypripedium
insigne_, in which there were two sepals and two petals. Of a similar
character was the flower found by Mr. J. A. Paine, and described in the
following terms by Professor Asa Gray in the 'American Journal of
Science,' July, 1866:--"The plant" (_Cypripedium candidum_) "bears two
flowers: the axillary one is normal; the terminal one exhibits the
following peculiarities. The lower part of the bract forms a sheath
which encloses the ovary. The labellum is wanting; and there are two
sterile stamens, the supernumerary one being opposite the other, _i.e._
on the side of the style where the labellum belongs. Accordingly the
first impression would be that the labellum is here transformed into a
sterile stamen. The latter, however, agrees with the normal sterile
stamen in its insertion as well as in shape, being equally adnate to the
base of the style. Moreover, the anteposed sepal is exactly like the
other, has a good midrib and an entire point. As the two sterile stamens
are anteposed to the two sepals, so are the two fertile stamens to the
two petals, and the latter are adnate to the style a little higher than
the former. The style is longer than usual, is straight and erect; the
broad, disciform stigma therefore faces upwards; it is oval and
symmetrical, and a light groove across its middle shows it to be
dimerous. The placentae, accordingly, are only two. The groove on the
stigma and the placentae are in line with the fertile stamens.
Here, therefore, is a symmetrical and complete, regular, but dimerous
orchideous flower, the first verticil of stamens not antheriferous, the
second antheriferous, the carpels alternate with these; and here we have
clear (and perhaps the first direct) demonstration that the orchideous
type of flower has two stamineal verticils, as Brown always insisted."
[Illustration: FIG. 198.--Regular dimerous flower of _Calanthe
vestita_.]
[Illustration: FIG. 199.--Regular dimerous flower of _Odontoglossum
Alexandrae_.]
Dr. Moore, of Glasnevin, kindly forwarded to the writer a flower of
_Calanthe vestita_ (fig. 198), in which there were two sepals only,
anterior and posterior, and two petals at right angles to the two
sepals. The lip was entirely wanting, but the column and ovary were in
their usual condition. In _Odontoglossum Alexandrae_ a similar reduction
of parts has been observed by the author (fig. 199).
It is curious to observe in these flowers how precisely one sepal
occupies the position of the labellum, and how the lateral petals are
displaced from the position they usually occupy, so as to form a regular
flower, the segments of which decussate, thus giving rise to a species
of regular peloria.
The genus _Maelenia_ was established on a malformed flower of _Orchis_ of
similar character to those above mentioned.
=Meiotaxy of the calyx.=--As already mentioned, this term is here
employed to denote those illustrations in which entire whorls are
suppressed. Complete deficiency of the calyx in a dichlamydeous flower
seems seldom or ever to occur; the nearest approach to it would be in
those cases where the calyx is, as it is termed, "obsolete," but here it
is chiefly the limb of the calyx which is atrophied, the lower portion
being more or less adherent to the ovary. In what are termed
monochlamydeous flowers both calyx and corolla are wanting, as in
_Salicineae_ and many other orders.
=Meiotaxy of the corolla.=--Deficiency of the entire corolla occurs in
conjunction with similar reductions in other organs, or as an isolated
phenomenon in the many apetalous varieties of plants recorded in books.
Deficiency of the corolla was observed in _Campanula perfoliata_ and
_Ruellia clandestina_ by Linne, who calls such blooms _flores
mutilati_.[480] Drs. Hooker and Thomson relate a similar occurrence in
_Campanula canescens_ and _C. colorata_. Some plants seem as a normal
occurrence to produce flowers of different construction, and are hence
termed dimorphic, as in many _Malpighiaceae_, _Violaceae_, _Oxalidaceae_,
in some of the flowers of which the petals are altogether wanting, while
in others the corolla is developed as usual. This deficiency of the
corolla is frequently, but not invariably, associated with an increased
fertility. Thus, in some violets the flowers produced in summer, and in
which the petals are either entirely suppressed or are more or less
atrophied, are always fertile, while the blossoms developed in spring,
and in which the petals are always present, are much less fertile. In
_Oxalis Acetosella_ there are two forms of flower, the one with, the
other without, petals, but both seem equally fertile. Linne remarks that
many plants which, in warm latitudes, produce a corolla, do not do so
when grown in colder climates. Thus, certain species of _Helianthemum_
are apetalous in Lapland. In the Pyrenees, according to Bentham, the
flowers of _Ajuga iva_ are constantly deprived of their corolla.[481]
Apetalous flowers have been noted most frequently in the following
plants:
Aconitum, sp. pl.!
Cardamine impatiens.
Cheiranthus Cheiri!
Viola odorata!
Cerastium vulgatum!
Alsine media.
Stellaria.
Lychnis dioica!
Dianthus barbatus, and other Caryophylleae.
Helianthemum, sp.!
Oxalis Acetosella.
Balsamineae.
Malpighiaceae.
Rosa centifolia.
arvensis!
Crataegus!
Medicago lupulina.
Melilotus officinalis.
Ononis minutissima.
Saxifraga longifolia.
Verbascum Thapsus.
Ajuga iva.
Teucrium Botrys.
Lamium purpureum!
amplexicaule.
Polemonium caeruleum.
Campanula, sp. pl.!
Ruellia clandestina.
Lonicera Periclymenum!
Tradescantia, sp.!
Hymenocallis.
The following references apply some to apetalous and others to
dimorphic flowers, but it must be remembered that the latter
plants are not necessarily wanting in petals or stamens, &c.,
though the functional activity of the parts may be impaired:
A. de Jussien, 'Monogr. Malpigh.,' pp. 82, 334. Torrey, 'Fl.
New York,' i, p. 428. Hooker and Thomson, 'Journ. Linn. Soc.,'
ii, p. 7, Guillemin, 'Archiv. de Botan.,' i, p. 412. Michalet,
'Bull. Soc. Bot. Fr.,' vii. p. 465. Mueller, 'Bot. Zeit.,' 1857,
p. 729. 'Natural History Review,' July, 1862, p. 235.
=Meiotaxy of the androecium.=--Complete suppression of the stamens
occurs normally in the female flowers of unisexual plants, and, as an
accidental occurrence, is not very uncommon. _Erica Tetralix_ is one of
the plants in which this is said to happen. The variety _anandra_ is
said to have been known in France since 1635. Cornuti speaks of it in
his 'Enchiridion.' In 1860 M. du Parquet discovered it in peaty woods
near Nangis (Seine et Marne).
Many _Umbelliferae_, such as _Trinia vulgaris_, present a like
deficiency, while it is of common occurrence among _Rosaceae_ and
_Pomaceae_. In the latter group the St. Valery apple, so often referred
to, is an illustration. To obtain fruits from this variety it is
necessary to apply pollen from another flower, a proceeding made the
occasion of festivity and rejoicing by the villagers in some parts of
France. In some of the _Artemisias_, especially in _Artemisia
Tournefortiana_, all the florets have been noticed to be female, owing
to the suppression of the stamens, and this suppression is associated
with a change in the form of florets.[482] Mr. Moggridge has
communicated to the author flowers of _Thymus Serpyllum_ from a plant in
which all the stamens were deficient, the flower being otherwise normal.
M. Dupont has given a list of nineteen species of _Chenopodiaceae_ in
which female flowers are occasionally produced, owing to the entire
suppression of the staminal whorl.[483]
Flowers the subjects either of regular or irregular peloria, _q. v._,
are often destitute of some or all their stamens, _e.g._ _Calceolaria_,
_Linaria_, _Viola_, &c., while in cases of synanthy suppression of some
of the parts of the flower, and specially of the stamens, is of very
common occurrence.
Suppression of the androecium as a teratological occurrence has been
most frequently noticed in the following plants, omitting members of
those families whose floral construction is normally incomplete in the
majority of instances, and exclusive also of cases of substitution. See
also under Heterogamy.
Ranunculus Ficaria!
auricomus!
bulbosus!
Cruciferae, sp. pl.
Violaceae, sp. pl.
Honckenya peploides.
Stellaria.
Caryophyllaceae, sp. pl.
Malpighiaceae, sp. pl.
Tropaeolum majus!
Fragaria vesca!
Rubus, sp.
Pyrus Malus.
Agrimonia vulgaris.
Rosaceae, sp. pl.
Trifolium hybridum.
repens.
Umbelliferae, sp. pl.
Onagraceae, sp. pl.
Hippuris vulgaris.
Callitriche vernalis.
autumnalis.
Lonicera Periclymenum.
Erica Tetralix.
Thymus Serpyllum.
Calceolaria.
Compositae, sp. pl.
Chenopodiaceae, sp. pl.
Stratiotes aloides.
=Meiotaxy of the gynoecium.=--Complete suppression of the pistil is of
more frequent occurrence than that of the stamens, hence more flowers
become accidentally unisexual by suppression of the pistil than by
deficiency of the stamens.
In many _Umbelliferae_, e.g. _Torilis Anthriscus_, _Cicuta virosa_, the
central flowers are often male, owing to the suppression of the pistil.
In many double flowers, owing to the excessive multiplication of
petaloid stamens, the pistil is suppressed, in which cases it often
happens that the flower is depressed in the centre, as in some garden
varieties of _Ranunculus_. Schlechtendal, in describing a flower of
_Colchicum autumnale_, in which the perianth was virescent, says that,
although the stamens were present, the pistil was absent.
In proliferous flowers the pistil is often completely defective, its
place being occupied by the adventitious bud or axis.
As in other cases of like nature, suppression of the pistil is very
frequently consequent on fusion of flowers or other changes. Thus Morren
relates an instance of synanthy in the flowers of _Torenia scabra_,
accompanied by resorption or disappearance of some parts and spiral
torsion of others. The pistil was entirely absent in this instance.[484]
M. Gaetano Licopoli places on record an instance where the petals and
carpels of _Melianthus major_ were suppressed.[485]
On the whole, the pistil seems less subject to changes of this character
than the androecium.
Suppression of the pistil has been most frequently recorded in flowers
(normally bisexual) of--
Ranunculus!
Aconitium!
Delphinium!
Paeonia.
Caryophylleae!
Umbelliferae.
Trifolium repens.
hybridum.
Compositae, sp. pl.
Datura.
Torenia asiatica.
Colchicum autumnale.
=Suppression of ovules,--abortion of seeds.=--The two cases are taken
together, as the effects are similar, though it must be remembered that
in the one case the ovules at any rate have been formed, but their
development has been arrested, while in the other they have never
existed. The precise cause that has determined the absence of seed
cannot in all cases be ascertained in the adult condition, hence it is
convenient to treat the two phenomena under one head.
Many plants in other than their native climates either produce no fruit
at all, or the fruits that are produced are destitute of seed, _e.g._
_Musa_, _Artocarpus_, &c. Some of the cultivated varieties of the grape
and of the berberry produce no seeds.
Suppression or abortion of the seed is frequently associated with the
excessive development either in size or number of other portions of the
plant, or with an altered condition, as when carpels become foliaceous
and their margins detached. Hybridisation and cross fertilisation are
also well-known agents in diminishing the number and size of seeds.
=Meiotaxy of the parts of the flower in general.=--In the preceding
sections suppression has been considered as it affected individual
members of a whorl or separate whorls. It rarely happens, however, that
the suppression is limited in this way. More generally several of the
parts of the flower are simultaneously affected in the same manner.
A few illustrations are all that is necessary to give as to this point.
One of the most familiar instances is that of the cauliflower or
broccoli, where the common flower-stalk is inordinately thickened and
fleshy, while the corolla and inner parts of the flower are usually
entirely suppressed; the four sepals can, however, generally be
detected.
Maximowicz describes a _Stellaria_ (_Kraschenikovia_) in which the upper
flowers are male only, while the lower ones, which ultimately become
buried in the soil, have neither petals, stamens, nor styles, but the
walls of the capsule are fleshy, and enclose numerous seeds.[486]
Kirschleger[487] mentions a variety of _Lonicera Caprifolium_, which was
not only destitute of petals but of stamens also.
In some species of _Muscari_ and _Bellevalia_ the uppermost flowers of
the raceme show more or less complete suppression of almost all the part
of which the flower normally consists. In those cases where an imperfect
perianth exists, but in which the stamens and pistils are entirely
suppressed, Morren applies the term Cenanthy, [Greek: kenos], empty.
=Complete suppression of the flower.=--It is not necessary in this place
to allude to that deficient production of flowers characteristic of what
is termed by gardeners a "sky bloomer." In such plants often the
requisite conditions are not complied with, and the skill of the
gardener is shown in his attempt to discover and allow the plant to
avail itself of the necessary requirements. We need here only allude to
those instances in which provision is made for the production of
flowers, and yet they are not produced. A good illustration of this is
afforded by the feather-hyacinth, _Hyacinthus comosus_, in which the
flowers are almost entirely suppressed, while the pedicels are
inordinately increased in number, and their colour heightened. Something
similar occurs in several allied species, and in _Bowiea volubilis_. The
wig plant (_Rhus Cotinus_) affords another illustration of the same
thing. Some tendrils also owe their appearance to the absence of
flowers, being modified peduncles; proofs of this may frequently be met
with in the case of the vine.
In _Lamium album_ I have seen one of the verticillasters on one side of
the stem completely wanting, the adjacent leaf being, however, as fully
formed as usual.
=General remarks on suppression.=--On comparing together the various
whorls of the flower in reference to suppression, and, it may be added,
to atrophy, we find that these phenomena occur most rarely in the calyx,
more frequently in the corolla, and very often in the sexual
organs and seeds; hence it would seem as if the uppermost and most
central organs, those most subject to pressure and latest in date of
development--formed, that is, when the formative energies of the plant
are most liable to be exhausted--are the most prone to be suppressed or
arrested in their development. When the plants in which these
occurrences happen most frequently are compared together, it may be seen
that partial or entire suppression of the floral envelopes, calyx, and
corolla, is far more commonly met with in the polypetalous and
hypogynous groups than in the gamopetalous or epigynous series.
The orders in which suppression (speaking generally) occurs most often
as a teratological occurrence are the following:--_Ranunculaceae_,
_Cruciferae_, _Caryophyllaceae_, _Violaceae_, _Leguminosae_, _Onagraceae_,
_Jasminaceae_, _Orchidaceae_. It will be observed that these are all
orders wherein suppression of the whole or part of the outer floral
whorls takes place in certain genera as a constant occurrence.
Again, it may be remarked that many of these orders show a tendency
towards a regular diminution of the assumed normal number of their
parts; thus, among _Onagraceae_, _Circeia_ and _Lopezia_ may be referred
to, the former normally dimerous, the latter having only one perfect
petal. So in fuchsias, a very common deviation consists in a trimerous
and rarely a dimerous symmetry of the flower.
Although, if the absolute number of genera or orders be counted, there
appears to be little difference in the frequency of the occurrence of
suppression in irregular flowers as contrasted with regular flowers, yet
if the individual instances could be counted in the two groups
respectively it would be found that suppression is more common among
irregular than in regular flowers. Thus, the number of individual
instances of flowers in which the perianth is defective is comparatively
large among _Violaceae_, _Leguminosae_, and _Orchidaceae_. This statement
hardly admits of precise statistical proof; still, it is believed that
any observer who pays attention to the subject must come to the same
conclusion. This is but another illustration of the fact that conditions
which are abnormal in one plant constitute the natural arrangement in
others.
As to the suppressions that occur in the case of the sexual organs, and
the relations they bear to dimorphism, diclinism, &c., but little stress
has been laid on them in this place, because their chief interest is in
a physiological point of view, and is treated of in the writings of
Mohl, Sprengel, Darwin, Hildebrand, and others. All that need be said
here is, that teratology affords very numerous illustrations of those
intermediate conditions which are also found, under natural
circumstances, between the absolutely unisexual flowers, male or female,
and the structurally hermaphrodite ones. Rudimentary stamens or pistils
are of very common occurrence in monstrous flowers. See Chapter on
Heterogamy, &c.
FOOTNOTES:
[465] 'Rev. Hortic.,' 1866, p. 467.
[466] De Rochebrune, 'Bull. Soc. Bot. Fr.,' ix, p. 281. The author
points out seven grades between complete absence of petals and their
presence in the normal number in this plant. See also Gaudin, in 'Koch.
Fl. Helv.;' Koch. 'Synops. Fl. Germ.;' Cramer, 'Bildungsabweich,' p. 85.
[467] 'Bull. Acad. Belg.,' t. xix, part 1, p. 255.
[468] 'Bull. Bot.,' i, p. 7, tab. i, f. 7.
[469] See Gay, 'Ann. Sc. Nat.,' iii, p. 27.
[470] 'Ann. Sc. Nat.,' 4 ser., v, p. 305.
[471] Cramer, 'Bildungsabweich,' p. 90.
[472] See also Clos, 'Bull. Soc. Bot. Fr.,' xiii, p. 96, adnot.
[473] See Cramer, 'Bildungsabweich,' p. 7. Hildebrand, 'Bot. Zeit.,' xx,
1862, p. 209.
[474] See Hildebrand, 'Bot. Zeit.,' xx, 1862, p. 209.
[475] 'Bull. Soc. Bot. Fr.,' viii, p. 287.
[476] 'Bull. Soc. Bot. Fr.,' vol. viii, 1861, p. 152.
[477] Ibid., ix, p. 275.
[478] Ibid., 1861, vol. viii, p. 149.
[479] 'Lobelia,' p. 55.
[480] 'Phil. Bot.,' p. 119.
[481] 'Cat. Plant. Pyr,' p. 58.
[482] Moquin-Tandon, loc. cit., p. 328.
[483] For other instances see Chatin in 'Ann. Sc. Nat.,' 4 ser., vol. v,
p. 305.
[484] See also Morren. 'Bull. Acad. Belg.,' xv, Fuchsia, p. 67.
[485] Cited in 'Bull. Soc. Bot., France,' t. xiv ("Rev. Bibl."), p. 253.
[486] 'Primit. Flor. Amurens.' p. 57.
[487] 'Flora.' 1848. p. 484.
BOOK IV.
DEVIATIONS FROM THE ORDINARY SIZE AND CONSISTENCE OF ORGANS.
In the animal kingdom the entire adult organism, as well as each of its
separate parts, has certain dimensions, beyond which, under ordinary
circumstances, it does not pass, either in the one direction or the
other. It may not be easy or possible to state what the limits are, but,
practically, this inability to frame a precise limitation is productive
of no inconvenience. It is universally admitted that a certain animal
attains such and such dimensions, and that one organ has a certain
proportionate size as contrasted with another. The same rules hold good
in the case of plants, though in them it is vastly more difficult to
ascertain what may be called the normal dimensions or proportions.
Nevertheless observation and experience soon show what may be termed the
average size of each plant, and any disproportion between the several
organs is speedily detected.
When there is a general reduction in size throughout all the organs of a
plant, or throughout all the nutritive organs, stem, leaves, &c., and
the several portions participate in this diminished size, we have what
are generally termed "dwarf varieties," dwarf in comparison, that is,
with the ordinary condition of the plants; on the other hand, if the
entire plant, or, at least, if the whole of one set of organs be
increased in size beyond the recognised average, we have large
varieties, often qualified by such terms as _macrophylla_, _longifolia_,
_macrantha_, &c. &c. In all these cases either the entire plant or whole
series of organs are alike increased or diminished beyond average
limits; and such variations are often very constant, and are transmitted
by hereditary transmission. It may be supposed that such deviations may
have originated, in the first instance, either from excessive use, or
from disuse, or from the agency of certain conditions promoting or
checking growth, as the case may be; but whether or no, it is certain
that these variations often persist under different conditions, and that
they often retain their distinctive characters side by side with plants
presenting the normal average dimensions. In other cases the variations
in size are of a less general character, and affect certain organs of a
whorl in a relative manner, as, for instance, in the case of didynamous
or tetradynamous stamens, where two or four stamens are longer than
their fellows, the long or short stamens and styles of di- and
tri-morphic flowers, &c. These differences are sometimes connected with
the development of parts in succession, and not simultaneously.
Teratological deviations of size differ from those of which mention has
just been made chiefly in this, that they are more limited in their
manifestations. It is not, as a rule, the whole plant, or the whole
series of nutritive or of reproductive organs, that are affected, but it
is certain parts only; the alteration in size is more a relative change
than an absolute one.
For convenience sake the teratological alterations of size may be
divided into those which are the result of increased growth and those
which arise from diminished action. It will be seen, therefore, that in
these instances it is the bulk of the organs that is increased, not
their number; moreover, their development or metamorphosis is not
necessarily altered. In connection with increased size an alteration of
consistence is so frequent that the two phenomena are here taken
together. It will be borne in mind that the changes of consistence from
membranous to succulent or woody are very frequent in the ordinary
course of development. They may also occur as accidental phenomena, or
the normal conditions of any particular flower or fruit may be exactly
reversed, the usually succulent fruit becoming dry and capsular, and so
forth.
PART I.
HYPERTROPHY.
The term hypertrophy may serve as a general one to comprise all the
instances of excessive growth and increased size of organs, whether the
increase be general or in one direction merely. General hypertrophy is
more a variation than a deformity, unless indeed it be caused by insect
puncture or the presence of a fungus, in which case the excessive size
results from a diseased condition. For our present purpose hypertrophy
may be considered as it affects the axile or the foliar organs, and also
according to the way in which the increased size is manifested, as by
increased thickness or swelling--intumescence, or by augmented
length-elongation, by expansion or flattening, or, lastly, by the
formation of excrescences or outgrowths, which may be classed under the
head of luxuriance or enation.
As size must be considered in this place relatively, it is not possible
to lay down any precise line separating what are considered to be the
normal dimensions from those which are abnormal.
In practice no inconvenience will be found to accrue from this inability
to establish a fixed rule, and we may say that an hypertrophied organ is
one which, from some cause or other, attains dimensions which are not
habitual to the plant in its usual, healthy, well-formed state.
It will be seen that under this general head of hypertrophy, increase of
size, however brought about, is included; thus, not only increase in
length, but also in thickness; alterations of substance or consistence,
no less than of dimensions, are here grouped together. The alterations
of consistence resulting from an inordinate development of cellular,
fibrous, or ligneous tissue, are, of course, strictly homologous with
the similar changes which occur, under ordinary circumstances, during
the ripening of fruits or otherwise.
Hypertrophy, whatever form it may assume, may be so slight as not
perceptibly to interfere with the functions of the part affected, or it
may exist to such an extent as to impair the due exercise of its office.
It may affect any or all parts of the plant, and is generally coexistent
with, if not actually dependent on, some other malformation. Thus, the
inordinate growth of some parts is most generally attended by deficiency
in the size and number of others, as in the peripheral florets of
_Viburnum_ or _Hydrangea_, where the corollas are relatively very large,
and the stamens and pistils abortive.
CHAPTER I.
ENLARGEMENT.
A swollen or thickened condition (_renflement_) is usually the result of
a disproportionate formation of the cellular tissue as contrasted with
the woody framework of the plant. We see marked instances of it in
cultivated carrots and turnips, the normal condition of the roots or
root-stocks in these plants being one of considerable hardness and
toughness, and their form slender, tapering, and more or less branched.
The disproportionate development of cellular tissue is also seen in
tubers and bulbs, and in the swollen stems of such plants as
_Echinocactus_, _Adenium obesum_, some species of _Vitis_, &c. So, too,
the upper portion of the flower-stalk occasionally becomes much
dilated, so as ultimately to form a portion of the fruit. But it is not
necessary to give farther illustrations of this common tendency in some
organs to become hypertrophied. As a result of injury from insects or
fungi, galls and excrescences of various kinds are very common, but
their consideration lies beyond the scope of the present work.
[Illustration: FIG. 200.--_Pelargonium_, one branch of which was
hypertrophied.]
=Enlargement of axile organs.=--All the species of _Pelargonium_,
_Geranium_, _Mirabilis,_ as well as those of _Caryophylleae_ and other
orders, have tumid nodes as a normal occurrence. In the genus
_Pelargonium_ this swelling is sometimes not confined to the nodes, but
extends to the interspaces between them, _e.g._ _P. spinosum_. This
condition, which happens as a natural feature in the species just named,
may also occur as an exceptional thing in others. The author is indebted
to Dr. Sankey for a branch of _Pelargonium_ which was thus thickened,
the remaining branches not being in any way affected. The leaves on the
swollen branch were smaller than the others, and their stalks more
flattened. There was, in this instance, no trace of fungus or insect to
account for the swelling of a single branch, which might, therefore, be
due to bud-variation, perhaps to reversion to some ancestral form. The
repeated cross fertilisations to which Pelargoniums have been subjected
render this hypothesis not an improbable one.
As an accompaniment to a spiral torsion of the woody fibres, this
distension of the stem is frequently met with, as in _Valeriana_,
_Dipsacus,_ &c. (See Spiral Torsion.)
=Knaurs.=--On certain trees, such as the oak, the hornbeam, some species
of _Crataegus_, &c., hard woody lumps may occasionally be seen
projecting, varying greatly in size, from that of a pea to that of a
cocoa-nut. They are covered with bark, and consist in the interior of
very hard layers of wood disposed irregularly, so as to form objects of
beauty for cabinet-makers' purposes. From the frequent presence of small
atrophied leaf-buds on their surface, it would seem as if the structures
in question were shortened branches, in which the woody layers had
become inordinately developed, as if by compensation for the curtailment
in length.[488] The cause of their formation is not known, but it has
been ascertained that they are not due to insect agency. Knaurs may
occasionally be used for purposes of propagation, as in the case of the
"uovoli of the olive" and the "burrs" that are formed on some varieties
of apple, from which both roots and leaf-shoots are produced in
abundance.
A distinction must be drawn between those instances in which the
swelling is solid throughout from the excessive formation of cellular
tissue, and those wherein it is hollow from the more rapid growth of the
outer as contrasted with the inner portions. These latter cases might be
classed under the head of distension.
[Illustration: FIG. 201.--Formation of tubers or hypertrophied buds in
the axils of leaves in the potato.]
=Enlargement of the buds= may be seen in the case of bulbs and tubers.
Occasionally these organs are developed in the axils of leaves, when
their nature becomes apparent. A swollen bud or bulbil in this
situation is not uncommon in some cultivated tulips and lilies. The
presence of small tubers in the axils of the leaves in the potato, as
shown in fig. 201, is also not unfrequent.
[Illustration: FIG. 202.--Inflorescence of ash (_Fraxinus_), with
hypertrophied pedicels, flowers absent.]
=Enlargement of the flower-stalk.=--The cauliflower and broccoli afford
familiar illustrations of hypertrophy of the flower-stalk, accompanied
by a corresponding defective development of the flowers. In the case of
the ash the terminal pedicels occasionally become swollen and distorted,
while the flowers are completely deficient, as shown in the adjacent cut
(fig. 202).
In grapes a similar condition may occasionally be met with in which the
terminal pedicels become greatly swollen and fused into a solid mass. It
would seem probable that this change is due to insect puncture, or to
the effect of fungus growth at an early stage of development, but as to
this point there is at present no evidence.[489]
[Illustration: FIG. 203.--Monstrous pear, showing extension and
ramification of the succulent floral axis. The bases of the sepals are
also succulent.]
In the apple a dilatation of the flower-stalk below the ordinary fruit
may occasionally be observed, thus giving rise to the appearance of two
fruits superposed and separated one from the other by a constriction.
(See fig. 176, p. 327.) The lower swelling is entirely axial in these
cases, as no trace of carpels is to be seen. M. Carriere[490] mentions
an instance wherein from the base of one apple projected a second
smaller one, destitute of carpels, but surmounted by calyx-lobes as
usual. The direction of this supernumerary apple was the exact opposite
of that of the primary fruit.
[Illustration: FIG. 204.--Monstrous pear, showing extension and swelling
of axis, &c.]
In pears, quinces, and apples, a not uncommon deviation is one in which
the axis is prolonged beyond the ordinary fruit, like which it is much
swollen. Occasionally the axis is not only prolonged, but even ramifies,
the branches partaking of the succulent character of the ordinary pome.
Such instances are frequently classed under the head of prolification,
but they have in general no claim to be considered in this light, for
the reasons already given in the chapter relating to that subject. (See
p. 135.)[491]
A very curious illustration of hypertrophy of the flower-stalk is
recorded and figured by M. Carriere[492] in the cherry. The calyx in
these fruits was completely superior, the succulent portion of the fruit
being made up of the dilated extremity of the peduncle, and possibly in
part of the base of the calyx. The general appearance was thus that of a
crab-apple. There was no stone in the interior, but simply a rudimentary
kernel or seed.[493]
Moquin-Tandon records an instance in which the stamens of each
individual flower in the inflorescence of a vine were hypertrophied, the
sepals, petals, and other organs of the flower, being proportionately
diminished.[494]
In this place may also be mentioned the hypertrophied condition of the
placenta observed by Alphonse de Candolle in a species of _Solanum_, and
also in a species of _Melastoma_. Not only was the placenta unusually
large in these flowers, but it also protruded beyond the ovary.[495] A
similar state of things in _Lobelia_ and _Cuphea_ has already been
alluded to under the head of Alterations of Direction (p. 210).
The following singular growth in a tomato is described by the Rev. M.
J. Berkeley in the 'Gardeners' Chronicle' for 1866, p. 1217, and appears
to have been an extension of the placenta:--"On the first glance it
seemed as if an unusually large grape-stone had accidentally fallen on
the upper surface of the fruit, and was attached by the narrow base. The
process was, however, five lines long, and much narrowed below, besides
which, though it was pale green above, the base was coral-red, like the
tomato itself. It grew on a narrow and shallow crack on the surface of
the fruit, and was found below to communicate directly with a
fibro-vascular bundle, which entered into the composition of a portion
of the placenta. On making a vertical section, instead of being
succulent, as I expected, it was white and spongy within, with several
lacunae, and one or two irregular fibro-vascular bundles, with highly
developed spiral vessels threading the centre. These vessels, moreover,
were tinged with brown, as in many cases of diseased tissues. There was
not the slightest appearance of placentae or anything indicating an
abortive fruit. On closer examination the cuticle was found to consist
of thick-walled cells, exactly like those of the tomato, while the
spongy mass consisted of a similar tissue to the fleshy portion of the
fruit, but with far less wrinkled walls, and more indistinct
intercellular spaces. The most striking point, however, was the immense
quantity of very irregular and unequal starch-grains with which they
were gorged, which gave a peculiar sparkling appearance to them when
seen _en masse_. I am inclined to regard the body rather as an abortive
axis than an undeveloped fruit. In almost all, if not all, these cases
of abnormal growth, whether from leaves, petioles, fruit, or other
portions of the plant, we find an immediate connection with one or more
spiral vessels, which if not existent at first are developed sooner or
later. In the present case the connection of the fibro-vascular tissue
of the fruit and abnormal growth was plain enough, but whether it
existed when the body was first given off I am unable to say, as it was
fully developed when the fruit was brought to me."
=Enlargement of the leaves.=--Increase in the size or substance of
leaves takes places in several ways, and affects the whole or only
certain portions of them. The simplest form of this malformation is met
with in our cabbages, which, by the art of the gardener, have been made
to produce leaves of greater size and thickness than those which are
developed in the wild form. In such instances the whole substance of the
leaf is increased in bulk, and the increase affects the fibrous
framework of the leaves as well as the cellular portions, though the
exaggerated development of the latter is out of proportion to that of
the former.
In some species of _Podocarpus_ there may occasionally be seen at the
base of the branchlets a dozen or more fleshy scales, of a rose colour,
passing gradually into the ordinary leaves of the plant, and evidently
analogous to the three fleshy confluent bracts which surround the ripe
fruit.
In other instances, while the fibrous framework of the leaf retains its
usual degree of development, the cellular parenchyma is developed in
excess, and, if the increase is so arranged that the number of
superposed layers of the cellular tissue is not increased, or their
thickness exaggerated, then we get such leaves as those of the "kail,"
or of the "Savoys" leaves, which are technically called by descriptive
botanists "folia bullata." In such leaves the disc of the leaf, rather
than the margin, is increased and its surface is thrown up into little
conical projections, which are hollow on the under side.
But leaves may increase beyond their usual size without such grave
alterations of form as those to which allusion has just been made. It is
well known that if a tree be cut down and new shoots be sent out from
the stump, the leaves formed on these shoots very often greatly exceed
the ordinary ones in dimensions. Such cases as this hardly come under
the head of malformations. But where one part only of the leaf is
excessively developed, the other portion remaining in its ordinary
condition, there can be no hesitation in ranking the phenomenon as
teratological.
Thus, Moquin says that the median nerve may be prolonged beyond the
blade of the leaf in the form of a short strap or ribbon-like
excrescence, while, at other times, the lateral parts of the leaf are
subjected to undue development. He refers to a case cited by
Schlotterbec[496] in which each side of the leaves of a yellow "violier"
(wallflower) was dilated into a kind of projecting lobe on either side
of the true apex of the leaf, thus rendering it in appearance
three-lobed. M. Delavaud[497] puts on record a case of hypertrophy in
the leaves of the common elm, resulting in the formation of an
additional lobe and a return to the tricostate type. A leaf so affected
is stated to have presented the appearance of a fusion of two leaves.
(See also Multiplication of leaves, p. 353.)
The hypertrophied and coloured leaf of _Gesnera_ occupying the place of
the absent inflorescence has been previously alluded to under the head
of displacement (p. 88).
In some instances hypertrophy is the opposite of suppression; as in the
case previously mentioned, where the stipule in the inflorescence of a
pea, which is usually undeveloped and rudimentary, was developed in the
form of a leafy cup or pitcher.
Another instance of the development of parts usually suppressed, is
afforded by the bud-scales of _Magnolia fuscata_, which may sometimes be
found with small but perfect leaves projecting from them, the leaf in
this case being the lamina which is ordinarily abortive, while the
scales are the representatives of the stipules. This condition is said
by Hooker and Thomson ('Flora Indica,' p. 73) to be constant in
_Magnolia Campbelli_.
=Enlargement of the perianth, &c.=--One or all the segments of the
perianth may be subjected to hypertrophy; thus, the utricle of _Carex
vulpina_ may frequently be observed to attain four or five times its
usual size, the contained ovary remaining unaffected. This condition is
generally the result of insect puncture. The growth of parasitic fungi
will produce a similar result, as is often seen in the common shepherd's
purse, _Thlaspi bursa pastoris_, and other _Cruciferae_. The perianth of
_Rumex aquaticus_ has been also observed to be occasionally
hypertrophied in conjunction with a similar condition of the pistil and
with atrophy of the ovules.
Moquin relates having found flowers of _Salsola Kali_ and of
_Chenopodium murale_ in which some of the segments of the perianth were
five or six times larger than they should be.
[Illustration: FIG. 205.--Hypertrophy of the perianth in _Cocos
nucifera_.]
The adjoining woodcut represents a singular condition of some cocoa-nuts
in the Kew Museum, the appearance of which is due apparently to an
hypertrophied condition of the segments of the perianth, which have not
only increased in length as the central nut has ripened, but have
developed in their tissues that fibrous tissue which ordinarily is found
in the pericarp only. This view of the structure of these nuts is borne
out by the fact that, under normal circumstances, the base of the
perianth contains a considerable amount of fibrous material. In the
present case this has increased to such an extent that the fruit appears
surrounded by a double husk, by an inner one as usual, and by an outer
six-parted one.
It will be remembered that in some of the _Cinchonaceae_, e.g.
_Mussaenda_, _Pinckneya_, _Calycophyllum_, one or more of the calycine
lobes are normally dilated and petaloid, the others remaining small and
comparatively inconspicuous. Inequality in size is, indeed, a common
occurrence in the sepals of many natural orders--_Polygalaceae_,
_Leguminosae_, _Labiatae_, &c. The flowers of a rose are mentioned by
Moquin as having presented an enlargement of the calyx without any other
alterations in form. Schlechtendal has noticed the same thing in
_Papaver Rhoeas_, Reichenbach in _Campanula persicifolia_, and A. de
Candolle in _C. Rapunculus_. M. Brongniart also has recorded[498] a
remarkable variety of _Primula sinensis_ cultivated in the Jardin des
Plantes at Paris, wherein the calyx is enormously developed. MM.
Fournier and Bonnet have described flowers of _Rubus_ with hypertrophied
calyx in conjunction with atrophy and virescence of the petals and other
changes.[499]
The corolla may be hypertrophied in some cases, though the change is
more rare than in most other organs. Moquin-Tandon mentions as subject
to this anomaly species of _Galeopsis_, _Prunella_, _Scabiosa_, and
_Dipsacus_, and also mentions a remarkable variety of _Viola odorata_
cultivated in the neighbourhood of Toulouse. The same learned author
also alludes to the so-called double Composites, viz. those in which the
usually tubular florets of the disc assume the form and proportions of
those of the ray, but these are hardly cases of hypertrophy.
=Enlargement of the androecium.=--Dunal[500] alludes to a curious
instance in a species of _Verbascum_, the lower flowers of which had
hairy stamens as usual, but the filaments of the topmost flower were
quite destitute of hairs, and dilated like a flat ribbon.
Moquin relates having found in the neighbourhood of Toulouse a plant of
_Solanum Dulcamara_ in which all the upper flowers had two or three
stamens of larger dimensions than the others. This happens habitually in
_Solanum tridynamum_ and _S. Amazonicum_, and to a less extent in _S.
vespertilio_ and _S. cornutum_; also in some species of _Hyoscyamus_.
These cases show the close affinity between the _Solanaceae_ and the
_Scrophulariaceae_.
=Enlargement of the gynoecium.=--In some flowers which have become
accidentally female the pistil becomes unusually large, and even to such
an extent as to prevent the passage of the pollen. Moquin remarks having
seen this enlargement in the pistils of _Suaeda fruticosa_ and _Kochia
scoparia_. The flowers of these Chenopods, under these circumstances,
resemble the female flowers of some nettles. The styles of _Anemone_ are
also much enlarged as the result of cultivation, and from their petaloid
appearance resemble those of the _Iris_ (Goethe). MM. Seringe and
Heyland[501] have figured some anomalous flowers of _Diplotaxis
tenuifolia_ in which the pistil, more or less distended and deformed,
was considerably elongated below, so that it seemed to be borne upon a
long stalk, analogous to that of fruits of Capparids. Dr. Klinsman[502]
mentions an instance of a similar kind combined with hypertrophy of the
sepals and pistils; indeed, the alteration is not uncommon among
Crucifers. _Pyrethrum inodorum_ is very subject to hypertrophy. The
styles of its radial florets become elongated without any other
alteration; at the same time the small corollas become green, and show a
tendency to assume a foliaceous condition. Sometimes the hypertrophy
affects also the styles of the central florets, and these also become
enlarged to double or treble their usual dimensions.
Linne has remarked that the ovary of _Tragopogon_ sometimes assumes very
large dimensions, as also does the pappus. He mentions a double-flowered
variety, the ovaries of which become ten or twelve times larger than
ordinary. M. Clos[503] records an instance in _Rumex scutatus_ wherein
the pistil was hypertrophied or club-shaped, and open at the top, or in
other cases funnel-shaped, three-lobed at the summit, each lobe
terminated by a style. One of the most frequent causes tending to the
hypertrophy of the pistil is attributable to the puncture of insects;
thus, when the ovary of _Juncus articulatus_ is thus punctured, it
acquires a size two or three times larger than ordinary, becoming at the
same time sterile.[504]
Occasionally the enlargement may be due to a fusion or incorporation of
other elements; thus, M. Lemaire describes an instance in which the
style of _Sinningia purpurea_ was much larger than ordinary, tubular,
bearing three small lobes, and altogether bearing much resemblance to
the column or "gynosteme" of Orchids. This appearance was due to the
cohesion and intimate union of the styles with three abortive
stamens.[505]
=Enlargement of the fruit.=--Most cultivated fruits are in a state of
true hypertrophy. Girod de Chantrans, after many trials, succeeded in
producing a peculiar variety of pea with pods double the ordinary
size.[506] M. Clos[507] mentions a case wherein the carpels of
_Delphinium dictyocarpum_ were hypertrophied. The change in size may or
may not be attended by a difference in form; thus, in certain
_Leguminosae_, as _Medicago lupulina_, _Melilotus leucantha_, the carpels
are sometimes hypertrophied and elongated, so as to resemble a claw or
hook.[508]
The fruit of the common groundsel (_Senecio vulgaris_) is in its normal
condition two or three times shorter than the involucre, and cylindrical
for its whole length, but it frequently happens that the fruits become
as long as the involucre itself, and taper from the base upwards, so as
to become beaked. Under this head may also be mentioned the fleshy
bulbils that are found in the capsules of _Crinum_, _Amaryllis_, and
_Agave_. These are true seeds enormously dilated.[509] In these seeds
the outer coating becomes very thick and fleshy, and is traversed by
spiral vessels.
It is obvious that very important results in a practical point of view
may be and have been arrived at by cultivators availing themselves of
this tendency of plants to increase in dimensions under certain
circumstances. It is needless to do more than refer to the many fruits,
vegetables, and cereals, which have thus become enlarged and improved by
careful selection and rearing.
=Alterations of consistence= often accompany changes in size. The change
may be one whereby the tissues become unusually hardened, by the
excessive formation of secondary woody deposits, or softer and more
succulent than ordinary, from the formation of an inordinate amount of
loose cellular tissue. Generally speaking, the appearances presented in
such cases are not sufficiently striking to demand notice other than as
regards their size. One illustration, however, may be cited from its
singularity. This was the case of a dahlia, in which the centre of the
flower was occupied by a projecting knob as large as a walnut, brown in
colour, and very hard in texture. This knob was nothing but the enlarged
and indurated extremity of the common receptacle, destitute of the
scales and florets which usually spring from it. No insect-puncture
could be detected, and no other reason for this peculiarity could be
ascertained.
FOOTNOTES:
[488] On the subject of knaurs, the reader is referred to Trecul, 'Ann.
Sc. Nat.,' 3 ser., vol. xx, p. 65; Lindley, 'Theory of Horticulture;'
Rev. M. J. Berkeley, 'Gardeners' Chronicle,' 1855, p. 756.
[489] Jaeger, 'Flora.' 1860. p. 49, tab. i.
[490] 'Revue Horticole,' 1868, p. 110, figs. 12, 13.
[491] The reader may also refer for further information on the subject
of malformed pears to Irmisch. 'Flora,' 1858, p. 38, tab. i; Lindley,
'Theory of Horticulture'; Caspary, 'Bull. Soc. Bot. France,' vol. vi,
1859 (Rev. Bibl.), p. 235; Duhamel, 'Phys. Arbr.,' liv. iii, cap. 3. p.
393, fig. 308; Bonnet, 'Recherch. Us. feuilles,' tab. xxvi, fig. 2;
Moquin-Tandon, 'El. Ter. Veg.,' p. 384, &c. Some of the cases recorded
are, however, instances of true prolification.
[492] 'Revue Horticole' 1868, p. 310.
[493] The interest of this accident is great, as showing how an
habitually superior ovary may become inferior--a change so rare in its
occurrence that its existence has been denied, and thus forming a marked
contrast with the frequency with which the converse change of an
inferior ovary to a superior one, from want of union with the calyx or
from imperfect development of the peduncle, may be observed. It is also
interesting as showing how the peduncle may become swollen, and at the
same time how the woody deposit of the endocarp may, as if by
compensation, be deficient. And, again, the malformation is not without
significance in regard to the relationship between the drupaceous and
the pomaceous subdivisions of _Rosaceae_. The case would fitly be
included under alterations of position, but the sheets relating to that
subject were printed off before the publication of M. Carriere's notice.
[494] 'Bull. Soc. Bot. France,' 1860, vol. vii, p. 881.
[495] "Monstr. Veget.," in 'Neue Denkschrift.'
[496] "Sched. de Monst. Plant." in 'Act. Helvet.,' t. ii, pl. ii, f. 14.
[497] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 144.
[498] 'Ann. Sc. Nat.,' ser. 2, t. i, p. 308, pl. ix _c_, fig. 1.
[499] 'Bull. Soc. Bot. France,' 1862, t. ix, p. 37.
[500] 'Consid. org. Fleur.,' Montpell., 1829, 25, 26, pl. ii, f. 18 and
19.
[501] 'Bull. Bot.,' t. i, p. 7, tab. 1.
[502] 'Linnaea,' vol. x, p. 604, tab. 5.
[503] 'Mem. Acad. Sc. Toulouse,' 5 ser., vol. iii.
[504] 'Re. nosol. Veget.,' pp. 342.
[505] 'Illustr. Hortic.,' 1868, Misc., p. 62.
[506] 'Ann. Soc. Linn.,' Paris, t. i, p. 139.
[507] 'Mem. Acad. Toulouse,' t. 6, 1862.
[508] 'D. C. Prod.,' ii, pp. 172, 187.
[509] Richard, "Obs. sur les bulbilles des Crinum;" 'Ann. Sc. Nat.,' t.
ii, p. 12. pl. i, fig. 1, 2. See also A. Braun, "Memoire sur les graines
charnues des Amaryllidees," &c.; 'Ann. Sc. Nat.,' 1860, vol. xiv, p. 1,
tab. 1.
CHAPTER II.
ELONGATION.
The class of cases coming under this head are sufficiently indicated by
the name. There are many instances of this phenomenon occurring under
different conditions, which, though unusual, can hardly be called
abnormal, such, for instance, as the great lengthening of roots in their
search for water, the excessive elongation that takes place in plants
when grown at a distance from the light, in their endeavour to attain to
which they become, as gardeners phrase it, "drawn." A similar result is
brought about in forests or plantations, where long spars are required,
by allowing the trees to grow very close to each other, so as to prevent
the lateral extension of the branches. When plants grow in running
water their roots, stems, and sometimes their leaves, become excessively
elongated, as in _Ranunculus fluitans_, the flower-stalks of _Valisneria
spiralis_, &c. These are cases of variation rather than of malformation,
but are none the less curious, or sometimes perplexing; thus, Lapeyrouse
described, in his 'Supplement a la flore des Pyrenees,' p. 27, under
the name _Potamogeton bifolium_, a plant which Mr. Bentham subsequently
discovered to be nothing but a flowerless variety of _Vicia Faba_
distorted by its growth in water.[510]
=Elongation of the root.=--This, as already remarked, is more often a
variation than a malformation, and is usually due to the presence of
water at a distance necessitating growth at the extremities of the root,
or to the presence of some obstacle, such as a stone, to avoid which the
root elongates till it has passed the obstruction. Occasionally in
Crocus corms some of the fibrils may be met with much lengthened and
thickened, and invested with a fleshy sheath. It is not certain,
however, that these structures are roots; possibly, nay probably, they
may be processes from the stem thrust downwards into the soil, similar
to the formations already described in the tulip (p. 85, fig. 39).
=Elongation of the inflorescence.=--Under this heading it is necessary
to consider lengthening of the common rachis in the case of an aggregate
inflorescence, and lengthening of the individual flower-stalks, whether
they be solitary or portions of a multiple inflorescence. The two
phenomena may occur together, but they are quite as often independent
one of the other. Thus, among _Umbelliferae_ the umbels are occasionally
met with supported on unusually long stalks, while the pedicels of the
individual flowers may or not be increased in length; so also with some
of the Composites, or the heads of flowers of some _Leguminosae_,
_Trifolium repens_, &c. &c.
Another illustration of the sort is that recorded by M. Fournier,
wherein the usually umbellate inflorescence of _Pelargonium_ was,
through the lengthening of the main stalk, transformed into a raceme.
Among Composites a similar change may sometimes be met with.
MM. Clos and De Schoenefeld have recorded the existence of a variety of
the sweet chestnut (_Castanea_)in which the female catkins were as long,
and bore nearly as many flowers, as the male spikes. This is stated to
be of constant occurrence in some localities, and to be accompanied by a
diminished size of the fruits. A similar elongation has been observed in
the case of the walnut, catkins of which have been seen bearing thirty
to thirty-five large nuts.[511]
In the strobile of the hop, _Humulus Lupulus_, a like elongation may
sometimes be met with, generally in association with a more or less
leafy condition of some of the scales.
Of a similar character, but complicated with extrusion or eversion of an
ordinarily concave axis, is the fig described by Zuccarini,[512] and
from the appearances presented by which that author draws the inference
that the peculiar appearance of the fig is due to the formation of a
large number of small bracts blended together for the greater part of
their length, and accompanied by the suppression of the internodes, and
consequent shortening of the axis. In the monstrous fruit the axis is
prolonged, and forms a kind of raceme or catkin, surrounded at the base
by numerous bracts, as in many _Amentaceae_. (See p. 204, figs. 105,
106.)
A lengthening of the axis of the female strobiles of _Coniferae_ is not
of infrequent occurrence in _Cryptomeria japonica_, _Larie europaea_,
&c., and this is usually associated, as has been before stated, with a
leaf-like condition of the bracts, and sometimes even with the
development of leaf-bearing shoots in place of the scales. (See under
Prolification of Inflorescence and Phyllomorphy, and for references, p.
115.)
=Elongation of the secondary flower-stalks.=--In the previous section
the effect of elongation of the main rachis has been considered. A
corresponding deviation occurs in the peduncles or pedicels, and
sometimes alters the general character of the inflorescence very
considerably, converting a spike into a raceme, a raceme into a corymb,
a capitulum into an umbel, and so forth. A few such cases may here be
alluded to. Fig. 206 represents a specimen of _Ranunculus acris_, in
which the lower and lateral flower-stalks were not only increased in
number, but so much lengthened as to form a flat-topped inflorescence--a
corymbose cyme. In many leguminous plants, as in _Trifolium repens_,
_Lotus corniculatus_, &c., what is usually a compact spike, or head of
flowers, becomes a raceme from the elongation of the pedicels. In
_Umbelliferae_ a similar change occurs, by virtue of which sometimes the
umbels themselves, and at other times the florets, are raised on
unusually long stalks, as in _Angelica Razoulzii_, _Carum Carui_,
_Thysselinum palustre_.[513] In _Compositae_, when affected by an
analogous change, the capitulum assumes the appearance of a simple
umbel, as in _Hypochaeris radicata_, _Senecio vulgaris_, and other
plants.
[Illustration: FIG. 206.--Inflorescence of _Ranunculus acris_, with
secondary peduncles lengthened.]
In some of the double-flowered apples which have been previously alluded
to, the flower-stalk is inordinately long when compared with the
adjacent ones. Possibly in some of these cases the absence of the usual
swelling of the upper part of the peduncle may be connected with its
increased length. One of the most striking instances of lengthened
flower-stalk occurred in an apple flower, wherein there was no swelling
beneath the calyx, while the latter was represented by five perfect
stalked leaves.
=Elongation of the leaves.=--In the case of water plants this change
keeps pace with the corresponding growth of the stem, _e.g._
_Ranunculus fluitans_, and in terrestrial plants there are varieties
termed longifoliar, from the unusual length of the leaves. A similar
lengthening occurs in the involucral leaves of _Umbelliferae_ and
_Compositae_, changing very materially the general aspect of the
inflorescence. Occasionally, also, the leaf-lobes of parsley (_Apium
Petroselinum_) and other crested-leaved plants may be observed to lose
their ordinary wavy form, and to be lengthened into flat riband-like
segments, as shown in fig. 207.
The only further illustrations that it is requisite to give of such
changes in this place are those occurring in lobed or compounded leaves,
which, from a lengthening of the midrib or central stalk, convert a
digitate or palmate leaf into a pinnate one. In these instances the
lobes or leaflets become separated one from another by a kind of
apostasis. This change may be frequently seen in the horse-chestnut,
particularly in the young shoots formed after the trees have been pruned
or pollarded. In the adjoining cut the intermediate stages between a
palmate or digitate leaf to a pinnate one may be seen. The specimens
from which the drawing was made were taken from the same tree at the
same time.
[Illustration: FIG. 207.--Portion of leaf of parsley, showing the change
from short wavy to long flat leaf-lobes.]
In the white clover, _Trifolium repens_, a similar transition may often
be observed, as also in some species of _Potentilla_.[514]
=Elongation of the parts of the flower.=--The only circumstance that
needs especial mention under this section is the great lengthening that
sometimes takes place in the carpels, sometimes as a result of injury
from insects or fungus, at other times without assignable cause.
[Illustration: FIG. 208.--Leaves of horse-chestnut, _AEsculus_, showing
passage from digitate to pinnate leaves.]
In the case of inferior ovaries this lengthening is, perhaps, even more
common, as in _Umbelliferae_, _Compositae_, &c. The common groundsel
(_Senecio vulgaris_) is especially liable to this form of enlargement of
the pistil, either in association with a leafy condition of the pappus
or without any such change.
=Elongation of the thalamus, placenta, &c.=--In some plants, as in
_Magnolia_ or _Myosurus_, the thalamus becomes much elongated, and bears
the carpels disposed spirally around it. A similar lengthening occurs in
malformed flowers, usually in association with a similar change in the
lower or outer part of the flower, by virtue of which the whorls become
separated from each other (Apostasis). Elongation and protrusion of the
placenta have been already alluded to at p. 119, and also at p. 125. In
some of these cases the elongated placenta has taken the form of a
leaf-bearing shoot.[515]
=Apostasis.=--Engelmann made use of this term to express the separation
of parts one from another by the unusual elongation of the
internodes.[516] He drew a distinction between the separation of
individual organs one from the other, and the corresponding displacement
of whorls. The subject has already been, to a considerable degree,
treated of in these pages under the head of dialysis, displacement, and
prolification, and but little need here be added. With reference to the
distance between one whorl and another, it will be remembered that,
although in the majority of cases the floral whorls are packed closely
together, yet in other instances the floral axis becomes elongated, and
thus separates the whorls one from another, by structures such as the
gynophores, androphores, &c., of _Passifloreae_, _Caryophylleae_,
_Capparideae_, &c. &c.
A similar elongation of the thalamus, bringing about the separation of
the floral whorls, or of their constituent parts, is very commonly met
with in association with median prolification. Where the individual
floral elements are thus thrown out of their usual verticillate
arrangement, they naturally assume a spiral disposition, and are, in
some cases, united by their margins, so that a spiral sheet or tube is
formed, surrounding the axis. This frequently occurs in double flowers
of the Chinese primrose, _Primula sinensis_.
Engelmann[517] figures a case wherein the calyx of _Anagallis
phoenicea_ was separated by a rather long internode from the corolla,
and a like illustration in _Torilis Anthriscus_.
[Illustration: FIG. 209.--Flower of _Delphinium_, showing apostasis of
carpels, from lengthening of the thalamus, &c. (Cramer.)]
A frequent change in Crucifers is due to the formation of a long stalk
bearing the pod, and thus giving rise to the appearance met as a
constant occurrence in Capparids.
In _Tropaeolum majus_ a similar elevation of the pistil may occasionally
be seen.
The adjacent figure of a monstrous _Delphinium_ taken from Cramer
illustrates well the elongation of the floral axis and the apostasis of
the carpels. In this instance the axis is terminated by a second flower
(median prolification).
One of the best-marked illustrations of these changes occurs in a
permanent malformation of _Epilobium hirsutum_, specimens of which were
originally obtained from the late Professor Henslow. The several floral
parts are here, some virescent, others truly foliaceous, and each whorl
is separated from its neighbour by a rather long internode. In _Fuchsia_
and _Campanula_ a like change may occasionally be observed.
Engelmann, in addition to those previously mentioned, cites the
following plants as having manifested this change:
_*Convallaria majalis!_, _*Tulipa Gesneriana!_, _Veronica Chamaedrys_,
_Orobanche gracilis_, _Solanum Lycopersicum_, _Gentiana campestris_,
_Hypericum_, _Helleborus fetidus_, _Caltha palustris_, _Brassica
oleracea!_ and many _Rosaceae_, _Caryophylleae_, _Cruciferae_, and
_Ranunculaceae_. (See Dialysis, Median Prolification, &c.)
Apostasis of the sub-floral or involucral leaves is not of infrequent
occurrence in malformations affecting _Compositae_ and _Umbelliferae_. In
the following genera it has been observed with especial
frequency:--_Torilis Anthriscus_, _Eryngium_, _Athamanta Cervaria_,
_Leontodon_, _Tragopogon pratense!_, _Wedelia perfoliata!_ In garden
anemones, also, it is a common deviation.
FOOTNOTES:
[510] 'Cat. Plant.,' Lang., p. 113.
[511] 'Bull. Soc. Bot. France,' t. i, 1854, p. 173, and t. xiii, p. 96.
[512] 'Abhandl. Math. Phys. Class.,' Band. iv, Abhandl. i, tab. i.
[513] See Cramer, 'Bildungsabweich,' pp. 62-79, and Fleischer,
'Missbild, der Culturpflanzen.'
[514] Schlechtendal, 'Bot. Zeit.,' 1844, p. 457; 'Linnaea,' xi, p. 301,
xiv, p. 363; 'Bot. Zeit.,' 1856, p. 72; Masters, 'Rep. Brit. Assoc.,'
Manchester, 1861; Coultas, 'What may be learnt from a tree,' p. 118.
[515] For further details refer to the chapter on Displacements, p. 86.
[516] 'De Antholysi,' p. 42, Sec. 49.
[517] Loc. cit., tab. 2, f. 6.
CHAPTER III.
ENATION.
Under the above heading are included certain forms arising from excess
not of growth, but of development, and consisting in the formation of
supplementary lobes or excrescences from various organs.
The new formations are not due either to a repetition or to a partition
of any organ, but are out-growths from others previously formed.
In prolification and in multiplication the adventitious structures are
of independent origin. In fission the new developments grow
simultaneously with the older ones, of which, indeed, they are mere
repetitions. Moreover, in fission the supplementary lobes do not, in
general, project a plan different from that of the original structure,
at least in the first instance, though their direction may ultimately
become changed.
In enation the new growth projects from a previously formed organ after
it has attained to considerable size, or even after its ordinary
proportions have been attained, and it sprouts out from the beginning in
a plane which is at a considerable angle to that of the parent organ,
and it is sometimes of a different structure from it, and has different
functions to fulfil.
Many of the instances that occur of scales projecting from petals, as in
_Caryophylleae_, _Sapindaceae_, &c., the coronal filaments of
passion-flowers, the cup of _Narcissus_, the appendages that beset the
segments of the perianth in _Lilium lancifolium_, and other similar
growths, may be referred to a like process. In many cases this has been
proved by a study of the development of the flower, from which it
appears that the growths in question are developed subsequently to the
formation of the ordinary floral whorls. It is requisite, however, to be
cautious in pronouncing upon the exact nature of these bodies, in the
absence of a knowledge of their period and mode of formation. They may
be mere outgrowths from one or other of the customary whorls, or they
may represent abortive stamens or petals, &c. Where circumstances
prevent the course of development from being traced, something may be
inferred as to their real nature from their position in regard to the
other parts of the flower, from their anatomical structure, and from
analogy or comparison with like organs in other plants. The period of
their formation is, perhaps, of less importance than was at one time
supposed, since it is well ascertained that, in some cases, the
formation of the parts of the flower, _e.g._ the stamens of mallows,
follows a centrifugal rather than a centripetal order.
In the case of monstrous developments of this nature too much care can
hardly be exercised, and the observer should rarely venture on an
explanation of the nature of the case from the evidence afforded by the
monstrous growth apart from that to be derived from the study of the
development and organization of the normal flower and from analogous
formations in allied plants.
=Excrescences from axile organs--Warts.=--In a preceding paragraph the
formation of gnaurs has been alluded to. There are other outgrowths,
called warts, occasionally met with in trees, and which are more closely
connected with the central tissues of the stem, while at the same time
they are not provided with buds, in which two particulars warts differ
from gnaurs.
Excrescences of this kind often attain a very large size, and may be
seen on old elms and other trees, but, as their formation is probably
more pathological than teratological, no further notice of these
structures need here be given. No special notice need here be taken of
the tubercles on the roots of so many _Leguminosae_, nor of the peculiar
excrescences on the roots of _Taxodium distichum_, as these appear to be
normal formations. But it may be well to mention in this place an
anomalous development which occurs occasionally in _Ruscus aculeatus_,
and in which, from the upper surface of the ordinary flattened leaf-like
branch, projects at right angles a second similar branch, so that in
section the appearance would be like that of the inverted letter t;
thus, [Symbol: Inverted upper-case T].
=Enation from foliar organs--Leaves.=--The development of adventitious
lobes from leaves may take place either from their surfaces or their
margins. A few illustrations may be given of each. In cabbage leaves a
formation of adventitious laminae projecting at right angles from the
primary one may frequently be observed. In the instance figured (fig.
210) the new growths proceeded almost exclusively from the thick midrib,
which, in the figure, is shown cut through just above the base. Not only
is the ordinary semilunar band of vascular tissue to be here seen, but a
similar broken line of vessels exists on the upper side of the
leaf-stalk; thus the whole structure resembles that of a stem or a
branch as much as that of a true leaf.
[Illustration: FIG. 210.--Section through base of midrib of cabbage
leaf, showing supplementary laminae, &c.]
The development of secondary leaves from the surfaces of primary ones
(phyllomania, autophyllogeny) has already been alluded to at p. 355.
Some of the cases wherein a leaf seems to have a double lamina may be
alluded to here, though possibly they would more properly be referred to
fission. The appearance presented is as if four wings projected from the
midrib, so that a cross section would be nearly in the form of
[Symbol: )O( turned 90 degrees.]. In an orange leaf presenting this
appearance the lower surface of one lamina was, as usual, dull in
colour, while the upper surface of the subjacent lamina was likewise
dull; hence the impression might arise that this was an instance of the
adhesion of two leaves back to back, but the petioles were not twisted,
as they must have been had two leaves thus been united, and neither in
the petiole nor in the midrib was there the slightest indication of
fusion, the vascular bundles being arranged in a circular manner, not in
a horseshoe-like arrangement, as would have been the case had adhesion
taken place.[518] (See p. 33.)
Such leaves as those of the hedgehog holly, _Ilex Aquifolium_, var.
_feroae_, and, to a less extent, bullate leaves, may also be mentioned
here as illustrations of hypertrophy or enation.
[Illustration: FIG. 211.--_Nephrodium molle_. Ordinary frond and forked
and crested varieties of the same, the crest arising from the inordinate
development of the margins of the pinnules.]
When the increased development occurs at the margin of the leaves,
especially, the result is a wavy or crisped appearance, "folia undulata,
_vel_ crispa."[519] These conditions occur normally in such leaves as
those of _Rumex crispus_, _Malva crispa_, &c., and are developed to an
extreme degree in garden varieties of parsley, some kails, &c., as well
as in many ferns, but these are probably cases rather of fission than
enation as here understood.[520]
=Enation from the sepals.=--The basal lobes of the calyx in _Campanula
Medium_, under normal circumstances, may be referred to in illustration
of this occurrence, while the adventitious spurs on the calyx of some
monstrous flowers seem due also to a like cause. These have already been
alluded to at p. 315.
=Enation from the corolla.=--The instances of this are more frequent
than in the case of the calyx, and admit of classification according as
they occur in polypetalous or gamopetalous flowers, on the outer or
inner surface of the petals, &c. Under natural circumstances the
formation of scales, lobes, &c., from the petals, as in some
_Caryophylleae_, _Sapindaceae_, &c. &c., may be explained, as already
remarked, by this process, rather than by fission, chorisis, or by
substitution of petals for stamens, &c. Each case must, however, be
examined on its own merits, as it is not safe to decide upon the
arrangement of parts in one flower by simply referring to the analogy of
others. In the following illustrations the course of development has
not, in all cases, been observed, and hence the explanation here given
must be taken with some reserve; for should it prove that the
adventitious lobes, &c., are formed simultaneously with the ordinary
petals, the case will be one of chorisis rather than of enation, as here
understood. Again, it may be that the supernumerary organs really
represent petals or stamens in disguise, though this hypothesis demands
the further assumption (in order to account for the interference with
the law of alternation) that suppression of certain organs has taken
place.
Taking first those instances in which the supplementary petals appear on
the inner surface of the corolla, as being at once the most frequent,
and as presenting the closest analogy, with similar conformations, under
natural circumstances, certain double-flowered varieties of the Chinese
primrose, _Primula sinensis_, may be mentioned. In these flowers the
calyx is normal, the tube of the corolla is traversed by ten vascular
bundles, and the limb is divided into ten fimbriated lobes. About
halfway up the tube, on the inner surface, are given off five
supernumerary petals, opposite to as many lobes of the corolla. Some of
the supplementary petals have a stamen in front of them, in the same
relative position as in the normal flower. In some cases the back or
outer surface of the supplementary petal is turned towards the inner or
upper surface of the primary corolla, thus [Symbol: ((turned 90 degrees
cw]; while, in other instances, the front of the adventitious lobe is
directed towards the corresponding surface of the original petal, thus
[Symbol: () turned 90 degrees]. Whether these supernumerary petals are
formed by chorisis or by enation cannot, with certainty, be determined
without examining the early stages of development.
[Illustration: FIG. 212.--_Datura fastuosa._ True corolla turned back to
show the supernumerary corolla with the petal-like segments attached to
its outer surface (reduced).]
Of more interest are those instances where the adventitious growth is on
the outside of the corolla; thus in a garden azalea there was
intermediate between the calyx and the corolla, both of which were
normal, a series of five petalodes, alternating with the sepals, and,
therefore, opposite to the lobes of the corolla, and adherent with them
at the very base, though elsewhere detached. These petalodes were
concave on the surface looking towards the calyx, and were there
brightly coloured, while the tint of the opposite surface looking
towards the corolla was of a duller hue, corresponding with that of the
outside of the corolla-tube. This arrangement of the colour was thus
precisely similar to that which occurred in the four-winged leaves
already referred to at p. 446. In some flowers of _Datura fastuosa_ a
similar series of excrescences was observed; the calyx and the corolla
were normal within the latter, intervening between it and the stamens
was a second corolla produced by duplication, and adherent to the inner
surface of this latter were five stamens. So far there was nothing very
peculiar; it remains to say, however, that on the outer surface of the
second corolla were five petal-like lobes closely adherent to it below,
but partially detached above. The colour of the adventitious segments
was paler on the outside than on the inner surface, as in the corolla
itself. The position of the several parts was such that they were
opposite one to the other; hence, while the lobes of the inner corolla
were opposite to those of the outer one, the intermediate petalodes were
opposite to both; thus:
S S S S S
--------------------------
P P P P P
X X X X X
--------------------------
P P P P P
st st st st st
The X indicating the position of the petalodes.
[Illustration: FIG. 213.--Gloxinia, with supernumerary segments on the
outside of the true corolla.]
A still more singular case is that of a variety of the Gloxinia,
described originally by Professor Edouard Morren,[521] but which is now
becoming common in English gardens. When first observed these flowers
were observed to produce petaloid segments outside the ordinary corolla,
and partially adherent to (or rather, not completely separated from it)
much as in the azalea before mentioned, the outer surface being brightly
coloured, like the inner surface of the corolla in ordinary gloxinias.
Being encouraged and tended by gardeners, in course of time, instead of
a series of petalodes, more or less distinct from one another, a second
corolla or "catacorolla" was formed outside the primary one, so that a
hose in hose flower was produced, but, in this case, the supplementary
flower was formed on the outside and not within the ordinary corolla.
Moreover, the disposition of the colour was reversed, for in the
outermost corolla the richest hues were on the outer surface, while in
the inner or true corolla they were on the inside.
Professor Morren considers the adventitious petalodes as rudiments of so
many supplementary flowers, axillary to the calyx, and adnate to the
corolla; each lobe then would, in this view, represent an imperfect
flower, and the completed catacorolla would be formed of a series of
confluent flowers of this description. But this view involves the
assumption of the suppression of all the parts of the flower, except the
lobes in question.
[Illustration: FIG. 214.--"Catacorolla" of _Gloxinia_, formed from the
union of adventitious petalodes on the outside of the true corolla
(after Morren).]
The view here propounded that the lobes in question are enations from
the true petals, which become confluent, so as to form the catacorolla,
is surely more simple, involves no assumptions of suppression of parts;
and moreover, is borne out by the examination of some flowers, where the
production of these adventitious lobes from the outside of the minute
partially developed petals could be distinctly seen.
=Enation from the stamens.=--An illustration of this process occurred in
some double-flowered rhododendrons, which presented the following
arrangement of parts:--calyx and corolla normal; within the latter eight
petal-like stamens, forming a pseudo-corolla. The appearance presented
by the petaloid filaments and anthers was as if they were adnate to the
centre of the petals, but, on closer examination, it appeared that the
petaloid expansion to which the dilated filament was apparently
attached, was equally a part of the stamens; in other words, that the
filament was provided with four petal-like wings, two on each side
[Symbol: 00 topped by (turned cw 90 deg., an o above and another ( turned cw
above that]. This disposition was well seen in the anther, half of which
was, in some cases, petaloid like the filament; in fact, the inner wing
of the latter was directly continuous with the petal-like expansion from
the anther. A section through the latter showed, going from within
outwards, the cut edges of two perfect polliniferous lobes in the
centre; and on either side the petaloid wing representing the remaining
anther-lobe; outside these were the edges of the remaining wings, one on
each side. (See p. 290, fig. 155.)
=Enation from the carpels.=--The only instances of this that need be
referred to are such cases as those in which spur-like projections,
horns, tubercles, or winged expansions, are formed from the surface of
the ovary during the course of its development. The extraordinary
cornute oranges described and figured by Ferrari, Gallesio, and other
writers on the genus _Citrus_, may be mentioned under this head. A
similar formation occurs in the fruit of some species of _Solanum_. (See
p. 316.)
FOOTNOTES:
[518] It is desirable in this place to allude to a singular case of
fissiparous division of a leaf of _Prunus Laurocerasus_ described by
Prof. Alexander Dickson ('Seemann's Journ. Botany,' vol. v, 1867, p.
323), and which did not come under the writer's notice till after the
sheet relating to fission, p. 61, had been sent to press. Dr. Dickson
thus speaks of this abnormal leaf:--"The petiole (unchanged) supported
two laminae, placed back to back, and united by their midribs (_i.e._ not
separated) to within about an inch from their extremities, which were
perfectly free from each other. These laminae stood vertically, their
edges being directed towards and away from the axis; and as they were
placed back to back, the shining surfaces, corresponding in structure to
the normal upper leaf-surface, were directed laterally outwards. In the
axil of this abnormal leaf were two axillary buds. The existence of two
leaf-apices and two axillary buds shows that this was not due to an
accidental exuberance of development, but to fissiparous division,
which, had it been complete, would have resulted in the replacement of a
single leaf by two leaves. The arrangement in Prof. Dickson's leaf may
be thus represented: [Symbol: )OO( with X above]. The nature of the case
may be even better seen by comparison with the normal arrangement, which
would be [Symbol: (OX turned 90 degrees ccw], while in those cases where
the fission of the leaf occurs in the same plane as that of the primary
lamina, as where a leaf splits into two lobes at the apex, with a midrib
to each, the arrangement is as follows: [Symbol: OX turned 90 degrees
ccw, with 2 arcs below forming a half circle], the X in all cases
representing the position of the axis, the O that of the axillary bud,
and the [Symbol: ( turned 90 degrees ccw] that of the laminae."
[519] Linn., 'Phil. Bot.,' Sec. 274. The term "_crispa_" is surely
preferable to that of Re, "phyllorhyseme."
[520] See C. Morren, "Consid. sur les deformations," &c., in 'Bull.
Acad. Belg.,' 1852, tom, xix, part 3, p. 444; and as to ferns, see
Moore, 'Nature-Printed British Ferns,' 8vo ed., where numerous
illustrations are given.
[521] 'Bull. Acad. Belg.,' t. xix, p. 224, tab. i; and 'Gardeners'
Chronicle,' 1865, p. 865.
PART II.
ATROPHY.
Under the head of atrophy are included those cases wherein the organs
affected are actually present, but in a dwarfed and stunted condition as
compared with surrounding parts.
The diminished size is, in such instances, obviously due to a partial
development and to an arrest of growth at a certain stage, from the
operation of various causes, either external or inherent to the
organization itself. It may affect any part of the plant, and exists, in
very varying degree, in different instances, being sometimes so slight
in amount as not to preclude the exercise of the functions of the part;
while in others, the structure is so incomplete that the office cannot
be performed. These differences depend, of course, upon the stage of
development which the organ had reached when its growth was checked. For
practical purposes atrophy may be distinguished from suppression by the
fact that in the latter case a certain element of the flower or plant
which, under ordinary circumstances, is present, is entirely wanting,
while, in the former class, it exists but in a rudimentary condition.
Again, atrophy is to be separated from that general diminution in the
size of the whole plant or of distinct parts of that plant which is
comprised under the term "nanism." Thus the several dwarf varieties of
plants (var. _nanae_), or those in which the leaves or flowers are
smaller than usual (var. _parvifoliae_, v. _parviflorae_), are truly
regarded as variations, and not as malformations properly so called.
Atrophy is partial and special in its operation, nanism is general.
Under ordinary circumstances atrophy is exemplified by the presence of
rudimentary or imperfect organs, as, for instance, in _Pentstemon_,
_Scrophularia_, &c., where one stamen is atrophied.
For convenience sake atrophy may be divided into abortion and
degeneration, the first including cases where, from arrest of
development occurring at an early stage, organs are present; but in a
much smaller and more rudimentary condition than usual, their form and
general appearance, except so far as regards their dimensions, not being
materially altered. On the other hand, in cases of degeneration,
development is not entirely checked, but rather perverted, so that not
only the dimensions are lessened, but the form is altered.
CHAPTER I.
ABORTION.
The sense in which this term is here understood has been explained in
the preceding paragraph. It is only necessary to say further, that cases
of abortion are to be distinguished from those of suppression, on the
one hand, and those of degeneration on the other. In suppression there
is from the first an absolute deficiency of a particular organ. In
degeneration the part is present, but in a diminished and perverted
condition. In abortion it exists, but in a stunted and dwarfed, but not
otherwise permuted state.
=Abortion of axile organs.=--When the main stem is arrested in its
growth, the habit and general appearance of the plant are materially
altered, as in the so-called stemless plants, _plantae acaules_. In these
the internodes are so slightly developed that the leaves are closely
crowded in tufts or rosettes. When this shortening of the stem
(acaulosia) occurs, without other considerable change in other organs,
the deviation is classed under the head of variation rather than of
monstrosity; and, indeed, in very many plants, this arrested growth of
the axis is the rule rather than the exception. When occurring in an
abnormal manner, atrophy of the stem is most frequently attended by
other more or less grave alterations in other structures; thus
Moquin-Tandon[522] cites an instance of _Camphorosma monspeliaca_,
wherein the stems presented the form of very short, hard, woody
tubercles, thickly clothed with deformed leaves, and invested by a vast
number of hairs, longer and more dense than usual. A similar deformity
sometimes occurs in an Indian species of _Artabotrys_; in these
specimens the branchlets are contracted in length, and bear numerous
closely packed scaly leaves, densely hairy, and much smaller than
ordinary.
Spines and thorns may he looked on as atrophied branches, and seem to
result from poorness of soil, as the same plants, which, in hungry land,
produce spines, develop their branches to the full extent when grown
under more favorable conditions.[523]
In the birch an arrest of development in some of the branches is of
common occurrence. The branch suddenly ceases to grow in length; at the
same time it thickens at the end into a large bulbous knob, from which
are developed a profusion of small twigs, whose direction is sometimes
exactly the reverse of that of the main branch. (See p. 347.)
The branches of the common spruce fir, especially the lateral ones, when
attacked by a particular species of aphis, are very apt to be developed
into a cone-like excrescence.[524]
A shortened condition of the flower-stalks occurs occasionally, greatly
altering the general character of the inflorescence. This has been
observed in pelargoniums and in the Chinese primrose, in both of which
the effect was to replace the umbellate form of inflorescence by a
capitate one.
=Abortion of the receptacle.=--Here may be mentioned those cases of
flowers with habitually inferior ovary (real or apparent), in which the
receptacle fails, from some cause or other, to dilate as usual. This has
already been alluded to under the head of Prolification, Displacements,
&c. (pp. 78, 130, &c., figs. 35-37, 64, &c.), and hence requires only
incidental comment in this place. There are, however, certain other
cases of a similar nature which may here be referred to; such as the
abortive condition of the inferior ovary, or rather of the receptacle,
that usually encircles the ovary in _Compositae_ and _Umbelliferae_. In
the former natural order the following plants have been met with in this
condition:--_*Tragopogon pratense!_, *_Cirsium arvense_, _Hypochaeris
radicata_, _Senecio vulgaris!_, _Coreopsis Drummondi_. In the latter
order, _Daucus Carota!_ _OEnanthe crocata!_ and _Thysselinum
palustre_, seem most frequently to have been observed in this
state.[525] In some gourds the receptacle may be seen partially
developed only, and forming a kind of cup, from which the true carpels
protrude.
=Abortion of the leaves.=--Arrest of growth in the leaves occurs in
different ways; sometimes the whole leaf is smaller than usual; at other
times certain parts only are reduced in size; while, in a third class of
cases, portions of the leaf are entirely suppressed.
Moquin[526] mentions having seen the leaves of _Chenopodium vulvaria_,
and of _Diplotaxis muralis_ reduced to a fourth of their natural size;
and he alludes to other cases of the same nature, seen by other
observers, in _Hypericum perforatum_ and _Blitum polymorphum_.
_Nicandra physaloides_[527] has also been met with in a similar
condition, which, indeed, is a common result of insect-puncture, and of
fungous growth in plants. Those instances in which the leaf is
diminished in size, without any attendant malformation in other organs,
may be regarded rather as variations than as monstrosities, as in the
case of the entire-leaved varieties of those plants which ordinarily
have cut or divided leaves, _e.g._ _Plantago Coronopus_, var.
_integrifolia_, _Papaver Rhoeas integrifolia_, &c. &c. The same remark
may be made of those specimens in which one part of the leaf is
developed to a less extent than another, as happens in the submerged
leaves of such plants as _Ranunculus aquatilis_, _Cabomba aquatica_, the
spiney leaves of _Berberis_, the fenestrated leaves of _Ouvirandra_, &c.
In the illustrations last cited the relative deficiency of one portion,
as contrasted with another, takes place as a constant occurrence, and is
uniform and regular throughout the whole leaf. When, on the other hand,
the deficiency in question happens accidentally and irregularly, the
change may be considered as a malformation. One side of the blade of the
leaf is frequently affected in this manner, the other portions remaining
unaffected. It would appear as if any plant might be thus altered, but
the following species appear to be particularly subject to this change:
_AEesculus Hippocastanum_, _Digitalis purpurea_, _Morus alba_, _Fagus
silvatica contracta_ (hort.), _Codiaeum variegatum_ var. _erosum_
(hort.), _Broussonettia papyrifera_, _Scolopendrium vulgare_, &c.
Frequently this irregular diminution in proportion is coexistent with an
unusual degree of cleavage or laciniation of the margin, as in _Acer
platanoides laciniatum_, _Tilia asplenifolia_, _Alnus imperialis_
(hort.), _Fagus silvatica_ var. (hort.), &c.