ON BUD-VARIATION, AND ON CERTAIN ANOMALOUS MODES OF REPRODUCTION AND
VARIATION.
BUD-VARIATIONS IN THE PEACH, PLUM, CHERRY, VINE, GOOSEBERRY, CURRANT,
AND BANANA, AS SHOWN BY THE MODIFIED FRUIT--IN FLOWERS: CAMELLIAS,
AZALEAS, CHRYSANTHEMUMS, ROSES, ETC.--ON THE RUNNING OF THE COLOUR IN
CARNATIONS--BUD-VARIATIONS IN LEAVES--VARIATIONS BY SUCKERS, TUBERS,
AND BULBS--ON THE BREAKING OF TULIPS--BUD-VARIATIONS GRADUATE INTO
CHANGES CONSEQUENT ON CHANGED CONDITIONS OF LIFE--CYTISUS ADAMI, ITS
ORIGIN AND TRANSFORMATION--ON THE UNION OF TWO DIFFERENT EMBRYOS IN ONE
SEED--THE TRIFACIAL ORANGE--ON REVERSION BY BUDS IN HYBRIDS AND
MONGRELS--ON THE PRODUCTION OF MODIFIED BUDS BY THE GRAFTING OF ONE
VARIETY OR SPECIES ON ANOTHER--ON THE DIRECT OR IMMEDIATE ACTION OF
FOREIGN POLLEN ON THE MOTHER-PLANT--ON THE EFFECTS IN FEMALE ANIMALS OF
A FIRST IMPREGNATION ON THE SUBSEQUENT OFFSPRING--CONCLUSION AND
SUMMARY.
This chapter will be chiefly devoted to a subject in many respects
important, namely, bud-variation. By this term I include all those sudden
changes in structure or appearance which occasionally occur in full-grown
plants in their flower-buds or leaf-buds. Gardeners call such changes
"Sports;" but this, as previously remarked, is an ill-defined expression,
as it has often been applied to strongly marked variations in seedling
plants. The difference between seminal and bud reproduction is not so great
as it at first appears; for each bud is in one sense a new and distinct
individual; but such individuals are produced through the formation of
various kinds of buds without the aid of any special apparatus, whilst
fertile seeds are produced by the concourse of the two sexual elements. The
modifications which arise through bud-variation can generally be propagated
to any extent by grafting, budding, cuttings, bulbs, &c., and occasionally
even by seed. Some few of our most beautiful and useful productions have
arisen by bud-variation.
Bud-variations have as yet been observed only in the vegetable {374}
kingdom; but it is probable that if compound animals, such as corals, &c.,
had been subjected to a long course of domestication, they would have
varied by buds; for they resemble plants in many respects. Thus any new or
peculiar character presented by a compound animal is propagated by budding,
as occurs with differently coloured Hydras, and as Mr. Gosse has shown to
be the case with a singular variety of a true coral. Varieties of the Hydra
have also been grafted on other varieties, and have retained their
character.
I will in the first place give all the cases of bud-variations which I have
been able to collect, and afterwards show their importance. These cases
prove that those authors who, like Pallas, attribute all variability to the
crossing either of distinct races, or of individuals belonging to the same
race but somewhat different from each other, are in error; as are those
authors who attribute all variability to the mere act of sexual union. Nor
can we account in all cases for the appearance through bud-variation of new
characters by the principle of reversion to long-lost characters. He who
wishes to judge how far the conditions of life directly cause each
particular variation ought to reflect well on the cases immediately to be
given. I will commence with bud-variations, as exhibited in the fruit, and
then pass on to flowers, and finally to leaves.
_Peach_ (_Amygdalus Persica_).--In the last chapter I gave two cases of
a peach-almond and double-flowered almond which suddenly produced fruit
closely resembling true peaches. I have also recorded many cases of
peach-trees producing buds, which, when developed into branches, have
yielded nectarines. We have seen that no less than six named and
several unnamed varieties of the peach have thus produced several
varieties of nectarine. I have shown that it is highly improbable that
all these peach-trees, some of which are old varieties, and have been
propagated by the million, are hybrids from the peach and nectarine,
and that it is opposed to all analogy to attribute the occasional
production of nectarines on peach-trees to the direct action of pollen
from some neighbouring nectarine-tree. Several of the cases are highly
remarkable, because, firstly, the fruit thus produced has sometimes
been in part a nectarine and in part a peach; secondly, because
nectarines thus suddenly produced have reproduced themselves by seed;
and thirdly, because nectarines are produced from peach-trees from seed
as well as from buds. The seed of the nectarine, on the other hand,
occasionally produces peaches; and we have seen in one instance that a
nectarine-tree yielded peaches by bud-variation. As the peach is
certainly the oldest or primary variety, the {375} production of
peaches from nectarines, either by seeds or buds, may perhaps be
considered as a case of reversion. Certain trees have also been
described as indifferently bearing peaches or nectarines, and this may
be considered as bud-variation carried to an extreme degree.
The _grosse mignonne_ peach at Montreuil produced "from a sporting
branch" the _grosse mignonne tardive_, "a most excellent variety,"
which ripens its fruit a fortnight later than the parent tree, and is
equally good.[813] This same peach has likewise produced by
bud-variation the _early grosse mignonne_. Hunt's large tawny nectarine
"originated from Hunt's small tawny nectarine, but not through seminal
reproduction."[814]
_Plums._--Mr. Knight states that a tree of the yellow magnum bonum
plum, forty years old, which had always borne ordinary fruit, produced
a branch which yielded red magnum bonums.[815] Mr. Rivers, of
Sawbridgeworth, informs me (Jan. 1863) that a single tree out of 400 or
500 trees of the Early Prolific plum, which is a purple kind, descended
from an old French variety bearing purple fruit, produced when about
ten years old bright yellow plums; these differed in no respect except
colour from those on the other trees, but were unlike any other known
kind of yellow plum.[816]
_Cherry_ (_Prunus cerasus_).--Mr. Knight has recorded (_idem_) the case
of a branch of a May-Duke cherry, which, though certainly never
grafted, always produced fruit, ripening later, and more oblong, than
the fruit on the other branches. Another account has been given of two
May-Duke cherry-trees in Scotland, with branches bearing oblong, and
very fine fruit, which invariably ripened, as in Knight's case, a
fortnight later than the other cherries.[817]
_Grapes_ (_Vitis vinifera_).--The black or purple Frontignan in one
case produced during two successive years (and no doubt permanently)
spurs which bore white Frontignan grapes. In another case, on the same
footstalk, the lower berries "were well-coloured black Frontignans;
those next the stalk were white, with the exception of one black and
one streaked berry;" and altogether there were fifteen black and twelve
white berries on the same stalk. In another kind of grape black and
amber-coloured berries were produced in the same cluster.[818] Count
Odart describes a variety which often bears on the same stalk small
round and large oblong berries; though the shape of the berry is
generally a fixed character.[819] Here is another striking case given
on the excellent authority of M. Carriere:[820] "a black Hamburgh grape
(Frankenthal) was cut down, and produced three suckers; one of these
was layered, and after a time produced much smaller berries, which
always ripened at least a fortnight {376} earlier than the others. Of
the remaining two suckers, one produced every year fine grapes, whilst
the other, although it set an abundance of fruit, matured only a few,
and these of inferior quality.
_Gooseberry_ (_Ribes grossularia_).--A remarkable case has been
described by Dr. Lindley[821] of a bush which bore at the same time no
less than four kinds of berries, namely, hairy and red,--smooth, small
and red,--green,--and yellow tinged with buff; the two latter kinds had
a different flavour from the red berries, and their seeds were coloured
red. Three twigs on this bush grew close together; the first bore three
yellow berries and one red; the second twig bore four yellow and one
red; and the third four red and one yellow. Mr. Laxton also informs me
that he has seen a Red Warrington gooseberry bearing both red and
yellow fruit on the same branch.
_Currant_ (_Ribes rubrum_).--A bush purchased as the Champagne, which
is a variety that bears blush-coloured fruit intermediate between red
and white, produced during fourteen years, on separate branches and
mingled on the same branch, berries of the red, white, and champagne
kinds.[822] The suspicion naturally arises that this variety may have
originated from a cross between a red and white variety, and that the
above transformation may be accounted for by reversion to both
parent-forms; but from the foregoing complex case of the gooseberry
this view is doubtful. In France, a branch of a red-currant bush, about
ten years old, produced near the summit five white berries, and lower
down, amongst the red berries, one berry half red and half white.[823]
Alexander Braun[824] also has often seen branches bearing red berries
on white currants.
_Pear_ (_Pyrus communis_).--Dureau de la Malle states that the flowers
on some trees of an ancient variety, the _doyenne galeux_, were
destroyed by frost: other flowers appeared in July, which produced six
pears; these exactly resembled in their skin and taste the fruit of a
distinct variety, the _gros doyenne blanc_, but in shape were like the
_bon-chretien_: it was not ascertained whether this new variety could
be propagated by budding or grafting. The same author grafted a
_bon-chretien_ on a quince, and it produced, besides its proper fruit,
an apparently new variety, of a peculiar form, with thick and rough
skin.[825]
_Apple_ (_Pyrus malus_).--In Canada, a tree of the variety called Pound
Sweet, produced,[826] between two of its proper fruit, an apple which
was well russetted, small in size, different in shape, and with a short
peduncle. As no russet apple grew anywhere near, this case apparently
cannot be accounted for by the direct action of foreign pollen. I shall
hereafter give {377} cases of apple-trees which regularly produce fruit
of two kinds, or half-and-half fruit; these trees are generally
supposed, and probably with truth, to be of crossed parentage, and that
the fruit reverts to both parent-forms.
_Banana_ (_Musa sapientium_).--Sir R. Schomburgk states that he saw in
St. Domingo a raceme on the Fig Banana which bore towards the base 125
fruits of the proper kind; and these were succeeded, as is usual,
higher up the raceme, by barren flowers, and these by 420 fruits,
having a widely different appearance, and ripening earlier than the
proper fruit. The abnormal fruit closely resembled, except in being
smaller, that of the _Musa Chinensis_ or _Cavendishii_, which has
generally been ranked as a distinct species.[827]
* * * * *
FLOWERS.--Many cases have been recorded of a whole plant, or single
branch, or bud, suddenly producing flowers different from the proper
type in colour, form, size, doubleness, or other character. Half the
flower, or a smaller segment, sometimes changes colour.
_Camellia._--The myrtle-leaved species (_C. myrtifolia_), and two or
three varieties of the common species, have been known to produce
hexagonal and imperfectly quadrangular flowers; and the branches
producing such flowers have been propagated by grafting.[828] The
Pompone variety often bears "four distinguishable kinds of
flowers,--the pure white and the red-eyed, which appear promiscuously;
the brindled pink and the rose-coloured, which may be kept separate
with tolerable certainty by grafting from the branches that bear them."
A branch, also, on an old tree of the rose-coloured variety has been
seen to "revert to the pure white colour, an occurrence less common
than the departure from it."[829]
_Crataegus oxycantha._--A dark pink hawthorn has been known to throw out
a single tuft of pure white blossoms;[830] and Mr. A. Clapham,
nurseryman, of Bradford, informs me that his father had a deep crimson
thorn grafted on a white thorn, which, during several years, always
bore, high above the graft, bunches of white, pink, and deep crimson
flowers.
_Azalea Indica_ is well known often to produce by buds new varieties. I
have myself seen several cases. A plant of _Azalea Indica variegata_
has been exhibited bearing a truss of flowers of _A. Ind. Gledstanesii_
"as true as could possibly be produced, thus evidencing the origin of
that fine variety." On another plant of _A. Ind. variegata_ a perfect
flower of _A. Ind. lateritia_ was produced; so that both _Gledstanesii_
and _lateritia_ no doubt originally appeared as sporting branches of
_A. Ind. variegata_.[831]
_Cistus tricuspis._--A seedling of this plant, when some years old,
produced, at Saharunpore,[832] some branches "which bore leaves and
flowers widely different from the normal form." "The abnormal leaf is
much less {378} divided, and not acuminated. The petals are
considerably larger, and quite entire. There is also in the fresh state
a conspicuous, large, oblong gland, full of a viscid secretion, on the
back of each of the calycine segments."
_Althaea rosea._--A double yellow Hollyock suddenly turned one year into
a pure white single kind; subsequently a branch bearing the original
double yellow flowers reappeared in the midst of the branches of the
single white kind.[833]
_Pelargonium._--These highly cultivated plants seem eminently liable to
bud-variation. I will give only a few well-marked cases. Gaertner has
seen[834] a plant of _P. zonale_ with a branch having white-edged
leaves, which remained constant for years, and bore flowers of a deeper
red than usual. Generally speaking, such branches present little or no
difference in their flowers: thus a writer[835] pinched off the leading
shoot of a seedling _P. zonale_, and it threw out three branches, which
differed in the size and colour of their leaves and stems; but on all
three branches "the flowers were identical," except in being largest in
the green-stemmed variety, and smallest in that with variegated
foliage: these three varieties were subsequently propagated and
distributed. Many branches, and some whole plants, of a variety called
_compactum_, which bears orange-scarlet flowers, have been seen to
produce pink flowers.[836] Hill's Hector, which is a pale red variety,
produced a branch with lilac flowers, and some trusses with both red
and lilac flowers. This apparently is a case of reversion, for Hill's
Hector was a seedling from a lilac variety.[837] Of all Pelargoniums,
Rollisson's Unique seems to be the most sportive; its origin is not
positively known, but is believed to be from a cross. Mr. Salter, of
Hammersmith, states[838] that he has himself known this purple variety
to produce the lilac, the rose-crimson or _conspicuum_, and the red or
_coccineum_ varieties; the latter has also produced the _rose d'amour_;
so that altogether four varieties have originated by bud variation from
Rollisson's Unique. Mr. Salter remarks that these four varieties "may
now be considered as fixed, although they occasionally produce flowers
of the original colour. This year _coccineum_ has pushed flowers of
three different colours, red, rose, and lilac, upon the same truss, and
upon other trusses are flowers half red and half lilac." Besides these
four varieties, two other scarlet Uniques are known to exist, both of
which occasionally produce lilac flowers identical with Rollisson's
Unique;[839] but one at least of these did not arise through
bud-variation, but is believed to be a seedling from Rollisson's
Unique.[840] There are, also, in the trade[841] two other slightly
different varieties, of unknown origin, of Rollisson's Unique: so that
altogether we have a curiously complex case {379} of variation both by
buds and seeds.[842] An English wild plant, the _Geranium pratense_,
when cultivated in a garden, has been seen to produce on the same plant
both blue and white, and striped blue and white flowers.[843]
_Chrysanthemum._--This plant frequently sports, both by its lateral
branches and occasionally by suckers. A seedling raised by Mr. Salter
has produced by bud-variation six distinct sorts, five different in
colour and one in foliage, all of which are now fixed.[844] The
varieties which were first introduced from China were so excessively
variable, "that it was extremely difficult to tell which was the
original colour of the variety, and which was the sport." The same
plant would produce one year only buff-coloured, and next year only
rose-coloured flowers; and then would change again, or produce at the
same time flowers of both colours. These fluctuating varieties are now
all lost, and, when a branch sports into a new variety, it can
generally be propagated and kept true; but, as Mr. Salter remarks,
"every sport should be thoroughly tested in different soils before it
can be really considered as fixed, as many have been known to run back
when planted in rich compost; but when sufficient care and time are
expended in proving, there will exist little danger of subsequent
disappointment." Mr. Salter informs me that with all the varieties the
commonest kind of bud-variation is the production of yellow flowers,
and, as this is the primordial colour, these cases may be attributed to
reversion. Mr. Salter has given me a list of seven differently coloured
chrysanthemums, which have all produced branches with yellow flowers;
but three of them have also sported into other colours. With any change
of colour in the flower, the foliage generally changes in a
corresponding manner in lightness or darkness.
Another Compositous plant, namely, _Centauria cyanus_, when cultivated
in a garden, not unfrequently produces on the same root flowers of four
different colours, viz., blue, white, dark-purple, and
particoloured.[845] The flowers of Anthemis also vary on the same
plant.[846]
_Roses._--Many varieties of the rose are known or are believed to have
originated by bud-variation.[847] The common double moss-rose was
imported into England from Italy about the year 1735.[848] Its origin
is unknown, but from analogy it probably arose from the Provence rose
(_R. centifolia_) by bud-variation; for branches of the common
moss-rose have several times been known to produce Provence roses,
wholly or partially destitute of moss: I have seen one such instance,
and several others have been recorded.[849] {380} Mr. Rivers also
informs me that he raised two or three roses of the Provence class from
seed of the old single moss-rose;[850] and this latter kind was
produced in 1807 by bud-variation from the common moss-rose. The white
moss-rose was also produced in 1788 by an offset from the common red
moss-rose: it was at first pale blush-coloured, but became white by
continued budding. On cutting down the shoots which had produced this
white moss-rose, two weak shoots were thrown up, and buds from these
yielded the beautiful striped moss-rose. The common moss-rose has
yielded by bud-variation, besides the old single red moss-rose, the old
scarlet semi-double moss-rose, and the sage-leaf moss-rose, which "has
a delicate shell-like form, and is of a beautiful blush colour; it is
now (1852) nearly extinct."[851] A white moss-rose has been seen to
bear a flower half white and half pink.[852] Although several
moss-roses have thus certainly arisen by bud-variation, the greater
number probably owe their origin to seed of moss-roses. For Mr. Rivers
informs me that his seedlings from the old single moss-rose almost
always produced moss-roses; and the old single moss-rose was, as we
have seen, the product by bud-variation of the double moss-rose
originally imported from Italy. That the original moss-rose was the
product of bud-variation is probable, from the facts above given and
from the moss-rose de Meaux (also a var. of _R. centifolia_)[853]
having appeared as a sporting branch on the common rose de Meaux.
Prof. Caspary has carefully described[854] the case of a six-year-old
white moss-rose, which sent up several suckers, one of which was
thorny, and produced red flowers, destitute of moss, exactly like those
of the Provence rose (_R. centifolia_): another shoot bore both kinds
of flowers and in addition longitudinally striped flowers. As this
white moss-rose had been grafted on the Provence rose, Prof. Caspary
attributes the above changes to the influence of the stock; but from
the facts already given, and from others to be given, bud-variation,
with reversion, is probably a sufficient explanation.
Many other instances could be added of roses varying by buds. The white
Provence rose apparently thus originated.[855] The double and
highly-coloured Belladonna rose has been known[856] to produce by
suckers both semi-double and almost single white roses; whilst suckers
from one of these semi-double white roses reverted to perfectly
characterised Belladonnas. Varieties of the China rose propagated by
cuttings in St. Domingo often revert after a year or two into the old
China rose.[857] Many cases {381} have been recorded of roses suddenly
becoming striped or changing their character by segments: some plants
of the Comtesse de Chabrillant, which is properly rose-coloured, were
exhibited in 1862,[858] with crimson flakes on a rose ground. I have
seen the Beauty of Billiard with a quarter and with half the flower
almost white. The Austrian bramble (_R. lutea_) not rarely[859]
produces branches with pure yellow flowers; and Prof. Henslow has seen
exactly half the flower of a pure yellow, and I have seen narrow yellow
streaks on a single petal, of which the rest was of the usual copper
colour.
The following cases are highly remarkable. Mr. Rivers, as I am informed
by him, possessed a new French rose with delicate smooth shoots, pale
glaucous-green leaves, and semi-double pale flesh-coloured flowers
striped with dark red; and on branches thus characterised there
suddenly appeared, in more than one instance, the famous old rose
called the Baronne Prevost, with its stout thorny shoots, and immense,
uniformly and richly coloured, double flowers; so that in this case the
shoots, leaves, and flowers, all at once changed their character by
bud-variation. According to M. Verlot[860] a variety called _Rosa
cannabifolia_, which has peculiarly shaped leaflets, and differs from
every member of the family in the leaves being opposite instead of
alternate, suddenly appeared on a plant of _R. alba_ in the gardens of
the Luxembourg. Lastly, "a running shoot" was observed by Mr. H.
Curtis[861] on the old Aimee Vibert Noisette, and he budded it on
Celine; thus a climbing Aimee Vibert was first produced and afterwards
propagated.
_Dianthus._--It is quite common with the Sweet William (_D. barbatus_)
to see differently coloured flowers on the same root; and I have
observed on the same truss four differently coloured and shaded
flowers. Carnations and pinks (_D. caryophyllus_, &c.) occasionally
vary by layers; and some kinds are so little certain in character that
they are called by floriculturists "catch-flowers."[862] Mr. Dickson
has ably discussed the "running" of particoloured or striped
carnations, and says it cannot be accounted for by the compost in which
they are grown: "layers from the same clean flower would come part of
them clean and part foul, even when subjected to precisely the same
treatment; and frequently one flower alone appears influenced by the
taint, the remainder coming perfectly clean."[863] This running of the
parti-coloured flowers apparently is a case of reversion by buds to the
original uniform tint of the species.
I will briefly mention some other cases of bud-variation to show how
many plants belonging to many orders have varied in their flowers;
numerous cases might be added. I have seen on a snap-dragon
(_Antirrhinum majus_) white, pink, and striped flowers on the same
plant, and branches with striped flowers on a red-coloured variety. On
a double stock (_Matthiola incana_) I have seen a branch bearing single
flowers; and {382} on a dingy-purple, double variety of the wall-flower
(_Cheiranthus cheiri_) a branch which had reverted to the ordinary
copper colour. On other branches of the same plant, some flowers were
exactly divided across the middle, one half being purple and the other
coppery; but some of the smaller petals towards the centre of these
same flowers were purple longitudinally streaked with coppery colour,
or coppery streaked with purple. A Cyclamen[864] has been observed to
bear white and pink flowers of two forms, the one resembling the
Persicum strain, and the other the Coum strain. _Oenothera biennis_ has
been seen[865] bearing flowers of three different colours. The hybrid
_Gladiolus colvillii_ occasionally bears uniformly coloured flowers,
and one case is recorded[866] of all the flowers on a plant thus
changing colour. A Fuchsia has been seen[867] bearing two kinds of
flowers. _Mirabilis jalapa_ is eminently sportive, sometimes bearing on
the same root pure red, yellow, and white flowers, and others striped
with various combinations of these three colours.[868] The plants of
the Mirabilis which bear such extraordinarily variable flowers, in
most, probably in all cases, owe their origin, as shown by Prof. Lecoq,
to crosses between differently-coloured varieties.
* * * * *
_Leaves and Shoots._--Changes, through bud-variation, in fruits and
flowers have hitherto been treated of, but incidentally some remarkable
modifications in the leaves and shoots of the rose and Cistus, and in a
lesser degree in the foliage of the Pelargonium and Chrysanthemum, have
been noticed. I will now add a few more cases of variation in
leaf-buds. Verlot[869] states that on _Aralia trifoliata_, which
properly has leaves with three leaflets, branches bearing simple leaves
of various forms frequently appear; these can be propagated by buds or
grafting, and have given rise, as he states, to several nominal
species.
With respect to trees, the history of but few of the many varieties
with curious or ornamental foliage is known; but several probably have
originated by bud-variation. Here is one case:--An old ash-tree
(_Fraxinus excelsior_) in the grounds of Necton, as Mr. Mason states,
"for many years has had one bough of a totally different character to
the rest of the tree, or of any other ash-tree which I have seen; being
short-jointed and densely covered with foliage." It was ascertained
that this variety could be propagated by grafts.[870] The varieties of
some trees with cut leaves, as the oak-leaved laburnum, the
parsley-leaved vine, and especially the fern-leaved beech, are apt to
revert by buds to the common form.[871] The fern-like leaves of the
beech sometimes revert only partially, and the branches display here
and there sprouts bearing common leaves, fern-like, and variously
shaped leaves. Such cases differ but little from the so-called {383}
heterophyllous varieties, in which the tree habitually bears leaves of
various forms; but it is probable that most heterophyllous trees have
originated as seedlings. There is a sub-variety of the weeping willow
with leaves rolled up into a spiral coil; and Mr. Masters states that a
tree of this kind kept true in his garden for twenty-five years, and
then threw out a single upright shoot bearing flat leaves.[872]
I have often noticed single twigs and branches on beech and other trees
with their leaves fully expanded before those on the other branches had
opened; and as there was nothing in their exposure or character to
account for this difference, I presume that they had appeared as
bud-variations, like the early and late fruit-maturing varieties of the
peach and nectarine.
Cryptogamic plants are liable to bud-variation, for fronds on the same
fern are often seen to display remarkable deviations of structure.
Spores, which are of the nature of buds, taken from such abnormal
fronds, reproduce, with remarkable fidelity, the same variety, after
passing through the sexual stage.[873]
With respect to colour, leaves often become by bud-variation zoned,
blotched, or spotted with white, yellow, and red; and this occasionally
occurs even with plants in a state of nature. Variegation, however,
appears still more frequently in plants produced from seed; even the
cotyledons or seed-leaves being thus affected.[874] There have been
endless disputes whether variegation should be considered as a disease.
In a future chapter we shall see that it is much influenced, both in
the case of seedlings and of mature plants, by the nature of the soil.
Plants which have become variegated as seedlings, generally transmit
their character by seed to a large proportion of their progeny; and Mr.
Salter has given me a list of eight genera in which this occurred.[875]
Sir F. Pollock has given me more precise information: he sowed seed
from a variegated plant of _Ballota nigra_ which was found growing
wild, and thirty per cent. of the seedlings were variegated; seed from
these latter being sown, sixty per cent. came up variegated. When
branches become variegated by bud-variation, and the variety is
attempted to be propagated by seed, the seedlings are rarely
variegated; Mr. Salter found this to be the case with plants belonging
to eleven genera, in which the greater number of the seedlings proved
to be green-leaved; yet a few were slightly variegated, or were quite
white, but none were worth keeping. Variegated plants, whether
originally produced from seeds or buds, can generally be propagated by
budding, grafting, &c.; but all are apt to revert by bud-variation to
their ordinary foliage. This tendency, however, differs much in the
varieties of even the same species; for instance, the golden-striped
variety of _Euonymus Japonicus_ "is very liable to run back to the
green-leaved, while the silver-striped {384} variety hardly ever
changes."[876] I have seen a variety of the holly, with its leaves
having a central yellow patch, which had everywhere partially reverted
to the ordinary foliage, so that on the same small branch there were
many twigs of both kinds. In the pelargonium, and in some other plants,
variegation is generally accompanied by some degree of dwarfing, as is
well exemplified in the "Dandy" pelargonium. When such dwarf varieties
sport back by buds or suckers to the ordinary foliage, the dwarfed
stature sometimes still remains.[877] It is remarkable that plants
propagated from branches which have reverted from variegated to plain
leaves[878] do not always (or never, as one observer asserts) perfectly
resemble the original plain-leaved plant from which the variegated
branch arose: it seems that a plant, in passing by bud-variation from
plain leaves to variegated, and back again from variegated to plain, is
generally in some degree affected so as to assume a slightly different
aspect.
* * * * *
_Bud-variation by Suckers, Tubers, and Bulbs._--All the cases hitherto
given of bud-variation in fruits, flowers, leaves, and shoots, have
been confined to buds on the stems or branches, with the exception of a
few cases incidentally noticed of varying suckers in the rose,
pelargonium, and chrysanthemum. I will now give a few instances of
variation in subterranean buds, that is, by suckers, tubers, and bulbs;
not that there is any essential difference between buds above and
beneath the ground. Mr. Salter informs me that two variegated varieties
of Phlox originated as suckers; but I should not have thought these
worth mentioning, had not Mr. Salter found, after repeated trials, that
he could not propagate them by "root-joints," whereas, the variegated
_Tussilago farfara_ can thus be safely propagated;[879] but this latter
plant may have originated as a variegated seedling, which would account
for its greater fixedness of character. The Barberry (_Berberis
vulgaris_) offers an analogous case; there is a well-known variety with
seedless fruit, which can be propagated by cuttings or layers; but
suckers always revert to the common form, which produces fruit
containing seeds.[880] My father repeatedly tried this experiment, and
always with the same result.
Turning now to tubers: in the common Potato (_Solanum tuberosum_) a
single bud or eye sometimes varies and produces a new variety; or,
occasionally, and this is a much more remarkable circumstance, all the
eyes in a tuber vary in the same manner and at the same time, so that
the whole tuber assumes a new character. For instance, a single eye in
a tuber of the {385} old _Forty-fold potato_, which is a purple
variety, was observed[881] to become white; this eye was cut out and
planted separately, and the kind has since been largely propagated.
_Kemp's Potato_ is properly white, but a plant in Lancashire produced
two tubers which were red, and two which were white; the red kind was
propagated in the usual manner by eyes, and kept true to its new
colour, and, being found a more productive variety, soon became widely
known under the name of _Taylor's Forty-fold_.[882] The _Old
Forty-fold_ potato, as already stated, is a purple variety; but a plant
long cultivated on the same ground produced, not as in the case above
given a single white eye, but a whole white tuber, which has since been
propagated and keeps true.[883] Several cases have been recorded of
large portions of whole rows of potatoes slightly changing their
character.[884]
Dahlias propagated by tubers under the hot climate of St. Domingo vary
much; Sir R. Schomburgk gives the case of the "Butterfly variety,"
which the second year produced on the same plant "double and single
flowers; here white petals edged with maroon; there of a uniform deep
maroon."[885] Mr. Bree also mentions a plant "which bore two different
kinds of self-coloured flowers, as well as a third kind which partook
of both colours beautifully intermixed."[886] Another case is described
of a dahlia with purple flowers which bore a white flower streaked with
purple.[887]
Considering how long and extensively many Bulbous plants have been
cultivated, and how numerous are the varieties produced from seed,
these plants have not varied so much by offsets,--that is, by the
production of new bulbs,--as might have been expected. With the
Hyacinth a case has been recorded of a blue variety which for three
successive years gave offsets which produced white flowers with a red
centre.[888] Another hyacinth has been described[889] as bearing on the
same truss a perfectly pink and a perfectly blue flower.
Mr. John Scott informs me that in 1862 _Imatophyllum miniatum_, in the
Botanic Gardens of Edinburgh, threw up a sucker which differed from the
normal form, in the leaves being two-ranked instead of four-ranked. The
leaves were also smaller, with the upper surface raised instead of
being channelled.
In the propagation of _Tulips_, seedlings are raised, called _selfs_ or
_breeders_, which "consist of one plain colour on a white or yellow
bottom. These, being cultivated on a dry and rather poor soil, become
broken or variegated and produce new varieties. The time that elapses
before they break varies from one to twenty years or more, and
sometimes this change never takes place."[890] The various broken or
variegated colours which give value to all tulips are due to
bud-variation; for although the {386} Bybloemens and some other kinds
have been raised from several distinct breeders, yet all the Baguets
are said to have come from a single breeder or seedling. This
bud-variation, in accordance with the views of MM. Vilmorin and
Verlot,[891] is probably an attempt to revert to that uniform colour
which is natural to the species. A tulip, however, which has already
become broken, when treated with too strong manure, is liable to flush
or lose by a second act of reversion its variegated colours. Some
kinds, as Imperatrix Florum, are much more liable than others to
flushing; and Mr. Dickson maintains[892] that this can no more be
accounted for than the variation of any other plant. He believes that
English growers, from care in choosing seed from broken flowers instead
of from plain flowers, have to a certain extent diminished the tendency
in flowers already broken to flushing or secondary reversion.
During two consecutive years all the early flowers in a bed of
_Tigridia conchiflora_[893] resembled those of the old _T. pavonia_;
but the later flowers assumed their proper colour of fine yellow
spotted with crimson. An apparently authentic account has been
published[894] of two forms of Hemerocallis, which have been
universally considered as distinct species, changing into each other;
for the roots of the large-flowered tawny _H. fulva_, being divided and
planted in a different soil and place, produced the small-flowered
yellow _H. flava_, as well as some intermediate forms. It is doubtful
whether such cases as these latter, as well as the "flushing" of broken
tulips and the "running" of particoloured carnations,--that is, their
more or less complete return to a uniform tint,--ought to be classed
under bud-variation, or ought to be retained for the chapter in which I
treat of the direct action of the conditions of life on organic beings.
These cases, however, have this much in common with bud-variation, that
the change is effected through buds and not through seminal
reproduction. But, on the other hand, there is this difference--that in
ordinary cases of bud-variation, one bud alone changes, whilst in the
foregoing cases all the buds on the same plant were modified together;
yet we have an intermediate case, for with the potato all the eyes in
one tuber alone simultaneously changed their character.
I will conclude with a few allied cases, which may be ranked either
under bud-variation, or under the direct action of the conditions of
life. When the common Hepatica is transplanted from its native woods,
the flowers change colour, even during the first year.[895] It is
notorious that the improved varieties of the Heartsease (_Viola
tricolor_) when transplanted often produce flowers widely different in
size, form, and colour: for instance, I transplanted a large
uniformly-coloured dark purple variety, whilst in full flower, and it
then produced much smaller, more elongated flowers, with the lower
petals yellow; these were succeeded by flowers marked with large purple
spots, and ultimately, towards the end of the same summer, by the
original large dark purple flowers. The slight changes which some {387}
fruit-trees undergo from being grafted and regrafted on various
stocks,[896] were considered by Andrew Knight[897] as closely allied to
"sporting branches," or bud-variations. Again, we have the case of
young fruit-trees changing their character as they grow old; seedling
pears, for instance, lose with age their spines and improve in the
flavour of their fruit. Weeping birch-trees, when grafted on the common
variety, do not acquire a perfect pendulous habit until they grow old:
on the other hand, I shall hereafter give the case of some weeping
ashes which slowly and gradually assumed an upright habit of growth.
All such changes, dependent on age, may be compared with the changes,
alluded to in the last chapter, which many trees naturally undergo; as
in the case of the Deodar and Cedar of Lebanon, which are unlike in
youth and closely resemble each other in old age; and as with certain
oaks, and with some varieties of the lime and hawthorn.[898]
Before giving a summary on Bud-variation I will discuss some singular and
anomalous cases, which are more or less closely related to this same
subject. I will begin with the famous case of Adam's laburnum or _Cytisus
Adami_, a form or hybrid intermediate between two very distinct species,
namely, _C. laburnum_ and _purpureus_, the common and purple laburnum; but
as this tree has often been described, I will be as brief as I can.
Throughout Europe, in different soils and under different climates,
branches on this tree have repeatedly and suddenly reverted to both
parent-species in their flowers and leaves. To behold mingled on the
same tree tufts of dingy-red, bright yellow, and purple flowers, borne
on branches having widely different leaves and manner of growth, is a
surprising sight. The same raceme sometimes bears two kinds of flowers;
and I have seen a single flower exactly divided into halves, one side
being bright yellow and the other purple; so that one half of the
standard-petal was yellow and of larger size, and the other half purple
and smaller. In another flower the whole corolla was bright yellow, but
exactly half the calyx was purple. In another, one of the dingy-red
wing-petals had a bright yellow narrow stripe on it; and lastly, in
another flower, one of the stamens, which had become slightly
foliaceous, was half yellow and half purple; so that the tendency to
segregation of character or reversion affects even single parts {388}
and organs.[899] The most remarkable fact about this tree is that in
its intermediate state, even when growing near both parent-species, it
is quite sterile; but when the flowers become pure yellow or pure
purple they yield seed. I believe that the pods from the yellow flowers
yield a full complement of seed; they certainly yield a large number.
Two seedlings raised by Mr. Herbert from such seed[900] exhibited a
purple tinge on the stalks of their flowers; but several seedlings
raised by myself resembled in every character the common laburnum, with
the exception that some of them had remarkably long racemes: these
seedlings were perfectly fertile. That such purity of character and
fertility should be suddenly reacquired from so hybridized and sterile
a form is an astonishing phenomenon. The branches with purple flowers
appear at first sight exactly to resemble those of _C. purpureus_; but
on careful comparison I found that they differed from the pure species
in the shoots being thicker, the leaves a little broader, and the
flowers slightly shorter, with the corolla and calyx less brightly
purple: the basal part of the standard-petal also plainly showed a
trace of the yellow stain. So that the flowers, at least in this
instance, had not perfectly recovered their true character; and in
accordance with this, they were not perfectly fertile, for many of the
pods contained no seed, some produced one, and very few contained as
many as two seeds; whilst numerous pods on a tree of the pure _C.
purpureus_ in my garden contained three, four, and five fine seeds. The
pollen, moreover, was very imperfect, a multitude of grains being small
and shrivelled; and this is a singular fact; for, as we shall
immediately see, the pollen-grains in the dingy-red and sterile flowers
on the parent-tree, were, in external appearance, in a much better
state, and included very few shrivelled grain. Although the pollen of
the reverted purple flowers was in so poor a condition, the ovules were
well-formed, and, when mature, germinated freely with me. Mr. Herbert
also raised plants from seeds of the reverted purple flowers, and they
differed _very little_ from the usual state of _C. purpureus_; but this
expression shows that they had not perfectly recovered their proper
character.
Prof. Caspary has examined the ovules of the dingy-red and sterile
flowers in several plants of _C. adami_ on the Continent,[901] and
finds them generally monstrous. In three plants examined by me in
England, the ovules were likewise monstrous, the nucleus varying much
in shape, and projecting irregularly beyond the proper coats. The
pollen-grains, on the other hand, judging from their external
appearance, were remarkably good, and readily protruded their tubes. By
repeatedly counting, under the microscope, the proportional number of
bad grains, Prof. Caspary ascertained that only 2.5 per cent. were bad,
which is a less proportion than in the pollen of three pure species of
Cytisus in their cultivated state, viz. _C. purpureus_, _laburnum_, and
_alpinus_. Although the pollen of _C. adami_ is thus in appearance
good, it does not follow, according {389} to M. Naudin's
observations[902] on Mirabilis, that it would be functionally
effective. The fact of the ovules of _C. adami_ being monstrous, and
the pollen apparently sound, is all the more remarkable, because it is
opposed to what usually occurs not only with most hybrids,[903] but
with two hybrids in the same genus, namely in _C. purpureo-elongatus_,
and _C. alpino-laburnum_. In both these hybrids, the ovules, as
observed by Prof. Caspary and myself, were well-formed, whilst many of
the pollen-grains were ill-formed; in the latter hybrid 20.3 per cent.,
and in the former no less than 84.8 per cent. of the grains were
ascertained by Prof. Caspary to be bad. This unusual condition of the
male and female reproductive elements in _C. adami_ has been used by
Prof. Caspary as an argument against this plant being considered as an
ordinary hybrid produced from seed; but we should remember that with
hybrids the ovules have not been examined nearly so frequently as the
pollen, and they may be much oftener imperfect than is generally
supposed. Dr. E. Bornet, of Antibes, informs me (through Mr. J.
Traherne Moggridge) that with hybrid Cisti the ovarium is frequently
deformed, the ovules being in some cases quite absent, and in other
cases incapable of fertilisation.
* * * * *
Several theories have been propounded to account for the origin of _C.
adami_, and for the transformations which it undergoes. These
transformations have been attributed by some authors to simple
bud-variation; but considering the wide difference between _C.
laburnum_ and _purpureus_, both of which are natural species, and
considering the sterility of the intermediate form, this view may be
summarily rejected. We shall presently see that, with hybrid plants,
two different embryos may be developed within the same seed and cohere;
and it has been supposed that _C. adami_ might have thus originated. It
is known that when a plant with variegated leaves is budded on a plain
stock, the latter is sometimes affected, and it is believed by some
that the laburnum has been thus affected. Thus Mr. Purser states[904]
that a common laburnum-tree in his garden, into which three _grafts_ of
the _Cytisus purpureus_ had been inserted, gradually assumed the
character of _C. adami_; but more evidence and copious details would be
requisite to make so extraordinary a statement credible.
Many authors maintain that _C. adami_ is a hybrid produced in the
common way by seed, and that it has reverted by buds to its two
parent-forms. Negative results are of little value; but Reisseck,
Caspary, and I myself, tried in vain to cross _C. laburnum_ and
_purpureus_; when I fertilised the former with pollen of the latter, I
had the nearest approach to success, for pods were formed, but in
sixteen days after the withering of the flowers they fell off.
Nevertheless, the belief that _C. adami_ is a spontaneously produced
hybrid between these two species is strongly supported by the fact that
hybrids between these species and two others have spontaneously {390}
arisen. In a bed of seedlings from _C. elongatus_, which grew near to
_C. purpureus,_ and was probably fertilised by it, through the agency
of insects (for these, as I know by experiment, play an important part
in the fertilisation of the laburnum), the sterile hybrid _C.
purpureo-elongatus_ appeared.[905] Thus, also, Waterer's laburnum, the
_C. alpino-laburnum_,[906] spontaneously appeared, as I am informed by
Mr. Waterer, in a bed of seedlings.
On the other hand, we have a clear and distinct account given by M.
Adam, who raised the plant, to Poiteau,[907] showing that _C. adami_ is
not an ordinary hybrid. M. Adam inserted in the usual manner a shield
of the bark of _C. purpureus_ into a stock of _C. laburnum_; and the
bud lay dormant, as often happens, for a year; the shield then produced
many buds and shoots, one of which grew more upright and vigorous with
larger leaves than the shoots of _C. purpureus_, and was consequently
propagated. Now it deserves especial notice that these plants were sold
by M. Adam, as a variety of _C. purpureus_, before they had flowered;
and the account was published by Poiteau after the plants had flowered,
but before they had exhibited their remarkable tendency to revert into
the two parent-species. So that there was no conceivable motive for
falsification, and it is difficult to see how there could have been any
error. If we admit as true M. Adam's account, we must admit the
extraordinary fact that two distinct species can unite by their
cellular tissue, and subsequently produce a plant bearing leaves and
sterile flowers intermediate in character between the scion and stock,
and producing buds liable to reversion; in short, resembling in every
important respect a hybrid formed in the ordinary way by seminal
reproduction. Such plants, if really thus formed, might be called
graft-hybrids.
* * * * *
I will now give all the facts which I have been able to collect
illustrative of the above theories, not for the sake of merely throwing
light on the origin of _C. adami_, but to show in how many
extraordinary and complex methods one kind of plant may affect another,
generally in connection with bud-variation. The supposition that either
_C. laburnum_ or _purpureus_ produced by ordinary bud-variation the
intermediate and the other form, may, as already remarked, be
absolutely excluded, from the want of any evidence, from the great
amount of change thus implied, {391} and from the sterility of the
intermediate form. Nevertheless such cases as nectarines suddenly
appearing on peach-trees, occasionally with the fruit half-and-half in
nature,--moss-roses appearing on other roses, with the flowers divided
into halves, or striped with different colours,--and other such cases,
are closely analogous in the result produced, though not in origin,
with the case of _C. adami_.
A distinguished botanist, Mr. G. H. Thwaites,[908] has recorded a
remarkable case of a seed from _Fuchsia coccinea_ fertilised by _F.
fulgens_, which contained two embryos, and was "a true vegetable twin."
The two plants produced from the two embryos were "extremely different
in appearance and character," though both resembled other hybrids of
the same parentage produced at the same time. These twin plants "were
closely coherent, below the two pairs of cotyledon-leaves, into a
single cylindrical stem, so that they had subsequently the appearance
of being branches on one trunk." Had the two united stems grown up to
their full height, instead of dying, a curiously mixed hybrid would
have been produced; but even if some of the buds had subsequently
reverted to both parent-forms, the case, although more complex, would
not have been strictly analogous with that of _C. adami_. On the other
hand, a mongrel melon described by Sageret[909] perhaps did thus
originate; for the two main branches, which arose from two
cotyledon-buds, produced very different fruit,--on the one branch like
that of the paternal variety, and on the other branch to a certain
extent like that of the maternal variety, the melon of China.
The famous _bizzarria Orange_ offers a strictly parallel case to that
of _Cytisus adami_. The gardener who in 1644 in Florence raised this
tree, declared that it was a seedling which had been grafted; and after
the graft had perished, the stock sprouted and produced the bizzarria.
Gallesio, who carefully examined several living specimens and compared
them with the description given by the original describer P. Nato,[910]
states that the tree produces at the same time leaves, flowers, and
fruit, identical with the bitter orange and with the citron of
Florence, and likewise compound fruit with the two kinds either blended
together, both externally and internally, or segregated in various
ways. This tree can be propagated by cuttings, and retains its
diversified character. The so-called trifacial orange of Alexandria and
Smyrna[911] resembles in its general nature the bizzarria, but differs
from it in the _sweet_ orange and citron being blended together in the
same fruit, and separately produced on the same tree: nothing is known
of its origin. In regard to the bizzarria, many authors believe that it
is a graft-hybrid; Gallesio on the other hand thinks that it is an
ordinary hybrid, with the habit of partially reverting {392} by buds to
the two parent-forms; and we have seen in the last chapter that the
species in this genus often cross spontaneously.
Here is another analogous, but doubtful case. A writer in the
'Gardener's Chronicle'[912] states that an _AEsculus rubicunda_ in his
garden yearly produced on one of its branches "spikes of pale yellow
flowers, smaller in size and somewhat similar in colour to those of _AE.
flava_." If as the editor believes _AEsculus rubicunda_ is a hybrid
descended on one side from _AE. flava_, we have a case of partial
reversion to one of the parent-forms. If, as some botanists maintain,
_AE. rubicunda_ is not a hybrid, but a natural species, the case is one
of simple bud-variation.
The following facts show that hybrids produced from seed in the
ordinary way, certainly sometimes revert by buds to their parent-forms.
Hybrids between _Tropaeolum minus_ and _majus_[913] at first produced
flowers intermediate in size, colour, and structure between their two
parents; but later in the season some of these plants produced flowers
in all respects like those of the mother-form, mingled with flowers
still retaining the usual intermediate condition. A hybrid Cereus
between _C. speciosissimus_ and _phyllanthus_,[914] plants which are
widely different in appearance, produced for the first three years
angular, five-sided stems, and then some flat stems like those of _C.
phyllanthus_. Koelreuter also gives cases of hybrid Lobelias and
Verbascums, which at first produced flowers of one colour, and later in
the season flowers of a different colour.[915] Naudin[916] raised forty
hybrids from _Datura laevis_ fertilised by _D. stramonium_; and three of
these hybrids produced many capsules, of which a half, or quarter, or
lesser segment was smooth and of small size like the capsule of the
pure _D. laevis_, the remaining part being spinose and of larger size
like the capsule of the pure _D. stramonium_: from one of these
composite capsules, plants were raised which perfectly resembled both
parent-forms.
Turning now to varieties. A _seedling_ apple, conjectured to be of
crossed parentage, has been described in France,[917] which bears
fruit, with one half larger than the other, of a red colour, acid
taste, and peculiar odour; the other side being greenish-yellow and
very sweet: it is said scarcely ever to include perfectly developed
seed. I suppose that this is not the same tree with that which
Gaudichaud[918] exhibited before the French Institute, bearing on the
same branch two distinct kinds of apples, one a _reinette rouge_, and
the other like a _reinette canada jaunatre_: this double-bearing
variety can be propagated by grafts, and continues to produce both
kinds; its origin is unknown. The Rev. J. D. La Touche sent me a
coloured drawing of an apple which he brought from Canada, of which
half, surrounding and including the whole of the calyx and the
insertion of the {393} footstalk, is green, the other half being brown
and of the nature of the _pomme gris_ apple, with the line of
separation between the two halves exactly defined. The tree was a
grafted one, and Mr. La Touche thinks that the branch which bore this
curious apple sprung from the point of junction of the graft and stock:
had this fact been ascertained, the case would probably have come into
the small class of graft-hybrids presently to be given. But the branch
may have sprung from the stock, which no doubt was a seedling.
Prof. H. Lecoq, who has made a great number of crosses between the
differently coloured varieties of _Mirabilis jalapa_,[919] finds that
in the seedlings the colours rarely combine, but form distinct stripes;
or half the flower is of one colour and half of a different colour.
Some varieties regularly bear flowers striped with yellow, white, and
red; but plants of such varieties occasionally produce on the same root
branches with uniformly coloured flowers of all three tints, and other
branches with half-and-half coloured flowers and others with marbled
flowers. Gallesio[920] crossed reciprocally white and red carnations,
and the seedlings were striped; but some of the striped plants also
bore entirely white and entirely red flowers. Some of these plants
produced one year red flowers alone, and in the following year striped
flowers; or conversely, some plants, after having borne for two or
three years striped flowers, would revert and bear exclusively red
flowers. It may be worth mentioning that I fertilised the _Purple
Sweet-pea_ (_Lathyrus odoratus_) with pollen from the light-coloured
_Painted Lady_: seedlings raised from one and the same pod were not
intermediate in character, but perfectly resembled both parents. Later
in the summer, the plants which had at first borne flowers identical
with those of the _Painted Lady_, produced flowers streaked and
blotched with purple; showing in these darker marks a tendency to
reversion to the mother-variety. Andrew Knight[921] fertilised two
white grapes with pollen of the Aleppo grape, which is darkly
variegated both in its leaves and fruit. The result was that the young
seedlings were not at first variegated, but all became variegated
during the succeeding summer; besides this, many produced on the same
plant bunches of grapes which were all black, or all white, or
lead-coloured striped with white, or white dotted with minute black
stripes; and grapes of all these shades could frequently be found on
the same footstalk.
In most of these cases of crossed varieties, and in some of the cases of
crossed species, the colours proper to both parents appeared in the
seedlings, as soon as they first flowered, in the form of stripes or larger
segments, or as whole flowers or fruit of two kinds borne on the same
plant; and in this case the appearance of the two colours cannot strictly
be said to be due to reversion, but to some incapacity of fusion, leading
to their {394} segregation. When, however, the later flowers or fruit,
produced during the same season or during a succeeding year or generation,
become striped or half-in-half, &c., the segregation of the two colours is
strictly a case of reversion by bud-variation. In a future chapter I shall
show that, with animals of crossed parentage, the same individual has been
known to change its character during growth, and to revert to one of its
parents which it did not at first resemble. From the various facts now
given there can be no doubt that the same individual plant, whether a
hybrid or a mongrel, sometimes returns in its leaves, flowers, and fruit,
either wholly or by segments, to both parent-forms, in the same manner as
the _Cytisus adami_, and the _Bizzarria Orange_.
* * * * *
We will now consider the few facts which have been recorded in support of
the belief that a variety when grafted or budded on another variety
sometimes affects the whole stock, or at the point of junction gives rise
to a bud, or graft-hybrid, which partakes of the characters of both stock
and scion.
It is notorious that when the variegated Jessamine is budded on the
common kind, the stock sometimes produces buds bearing variegated
leaves: Mr. Rivers, as he informs me, has seen instances of this. The
same thing occurs with the Oleander.[922] Mr. Rivers, on the authority
of a trustworthy friend, states that some buds of a golden-variegated
ash, which were inserted into common ashes, all died except one; but
the ash-stocks were affected,[923] and produced, both above and below
the points of insertion of the plates of bark bearing the dead buds,
shoots which bore variegated leaves. Mr. J. Anderson Henry has
communicated to me a nearly similar case: Mr. Brown, of Perth, observed
many years ago, in a Highland glen, an ash-tree with yellow leaves; and
buds taken from this tree were inserted into common ashes, which in
consequence were affected, and produced the _Blotched Breadalbane Ash_.
This variety has been propagated, and has preserved its character
during the last fifty years. Weeping ashes, also, were budded on the
affected stocks, and became similarly variegated. Many authors consider
variegation as the result of disease; and on this view, which however
is doubtful, for some variegated plants are perfectly healthy and
vigorous, the foregoing cases may be looked at as the direct result of
the inoculation of a disease. Variegation is much influenced, as we
shall hereafter see, by the nature of the soil in which the {395}
plants are grown; and it does not seem improbable that whatever change
in the sap or tissues certain soils induce, whether or not called a
disease, might spread from the inserted piece of bark to the stock. But
a change of this kind cannot be considered to be of the nature of a
graft-hybrid.
There is a variety of the hazel with dark-purple leaves, like those of
the copper-beech: no one has attributed this colour to disease, and it
apparently is only an exaggeration of a tint which may often be seen on
the leaves of the common hazel. When this variety is grafted on the
common hazel,[924] it sometimes colours, as has been asserted, the
leaves below the graft; but I should add that Mr. Rivers, who has
possessed hundreds of such grafted trees, has never seen an instance.
Gaertner[925] quotes two separate accounts of branches of dark and
white-fruited vines which had been united in various ways, such as
being split longitudinally, and then joined, &c.; and these branches
produced distinct bunches of grapes of the two colours, and other
bunches with grapes either striped or of an intermediate and new tint.
Even the leaves in one case were variegated. These facts are the more
remarkable because Andrew Knight never succeeded in raising variegated
grapes by fertilising white kinds by pollen of dark kinds; though, as
we have seen, he obtained seedlings with variegated fruit and leaves,
by fertilising a white variety by the variegated dark Aleppo grape.
Gaertner attributes the above-quoted cases merely to bud-variation; but
it is a strange coincidence that the branches which had been grafted in
a peculiar manner should alone have thus varied; and H. Adorne de
Tscharner positively asserts that he produced the described result more
than once, and could do so at will, by splitting and uniting the
branches in the manner described by him.
I should not have quoted the following case had not the author of 'Des
Jacinthes'[926] impressed me with the belief not only of his extensive
knowledge, but of his truthfulness: he says that bulbs of blue and red
hyacinths may be cut in two, and that they will grow together and throw
up a united stem (and this I have myself seen), with flowers of the two
colours on the opposite sides. But the remarkable point is, that
flowers are sometimes produced with the two colours blended together,
which makes the case closely analogous with that of the blended colours
of the grapes on the united vine-branches.
Mr. E. Trail stated in 1867, before the Botanical Society of Edinburgh
(and has since given me fuller information), that several years ago he
cut about sixty blue and white potatoes into halves through the eyes or
buds, and then carefully joined them, destroying at the same time the
other eyes. Some of these united tubers produced white, and others blue
tubers; and it is probable that in these cases the one half alone of
the bud grew. Some, however, produced tubers partly white and partly
blue; and the tubers from about four or five were regularly mottled
with the two colours. in these latter cases we may conclude that a stem
had been formed by {396} the union of the bisected buds; and as tubers
are produced by the enlargement of subterranean branches arising from
the main stem, their mottled colour apparently affords clear evidence
of the intimate commingling of the two varieties. I have repeated these
experiments on the potato and on the hyacinth on a large scale, but
with no success.
The most reliable instance known to me of the formation of a
graft-hybrid is one, recorded by Mr. Poynter,[927] who assures me, in a
letter of the entire accuracy of the statement, _Rosa Devoniensis_ had
been budded some years previously on a white Banksian rose; and from
the much enlarged point of junction, whence the Devoniensis and
Banksian still continued to grow, a third branch issued, which was
neither pure Banksian nor pure Devoniensis, but partook of the
character of both; the flowers resembled, but were superior in
character to those of the variety called _Lamarque_ (one of the
Noisettes), while the shoots were similar in their manner of growth to
those of the Banksian rose, with the exception that the longer and more
robust shoots were furnished with prickles. This rose was exhibited
before the Floral Committee of the Horticultural Society of London. Dr.
Lindley examined it, and concluded that it had certainly been produced
by the mingling of _R. Banksiae_ with some rose like _R. Devoniensis_,
"for while it was very greatly increased in vigour and in the size of
all the parts, the leaves were half-way between a Banksian and
Tea-scented rose." It appears that rose-growers were aware that the
Banksian rose sometimes affects other roses. Had it not been for this
latter statement, it might have been suspected that this new variety
was simply due to bud-variation, and that it had occurred by a mere
accident at the point of junction between the two old kinds.
To sum up the foregoing facts: the statement that _Cytisus adami_
originated as a graft-hybrid is so precise that it can hardly be rejected,
and, as we have just seen, some analogous facts render the statement to a
certain extent probable. The peculiar, monstrous condition of the ovules,
and the apparently sound condition of the pollen, favour the belief that it
is not an ordinary or seminal hybrid. On the other hand, the fact that the
same two species, viz. _C. laburnum_ and _purpureus_, have spontaneously
produced hybrids by seed, is a strong argument in support of the belief
that C. _adami_ originated in a similar manner. With respect to the
extraordinary tendency which this tree exhibits to complete or partial
reversion, we have seen that undoubted seminal hybrids and mongrels are
similarly liable. On the whole, I am inclined to put trust in M. Adam's
statement; and if it should ever be proved true, the same view would
probably have {397} to be extended to the Bizzarria and Trifacial oranges
and to the apples above described; but more evidence is requisite before
the possibility of the production of graft-hybrids can be fully admitted.
Although it is at present impossible to arrive at any certain conclusion
with respect to the origin of these remarkable trees, the various facts
above given appear to me to deserve attention under several points of view,
more especially as showing that the power of reversion is inherent in Buds.
* * * * *
_On the direct or immediate action of the Male Element on the Mother
Form._--Another remarkable class of facts must be here considered, because
they have been supposed to account for some cases of bud-variation: I refer
to the direct action of the male element, not in the ordinary way on the
ovules, but on certain parts of the female plant, or in the case of animals
on the subsequent progeny of the female by a second male. I may premise
that with plants the ovarium and the coats of the ovules are obviously
parts of the female, and it could not have been anticipated that they would
be affected by the pollen of a foreign variety or species, although the
development of the embryo, within the embryonic sack, within the ovule,
within the ovarium, of course depends on the male element.
Even as long ago as 1729 it was observed[928] that white and blue
varieties of the Pea, when planted near each other, mutually crossed,
no doubt through the agency of bees, and in the autumn blue and white
peas were found within the same pods. Wiegmann made an exactly similar
observation in the present century. The same result has followed
several times when a variety with peas of one colour has been
artificially crossed by a differently-coloured variety.[929] These
statements led Gaertner, who was highly sceptical on the subject,
carefully to try a long series of experiments: he selected the most
constant varieties, and the result conclusively showed that the colour
of the skin of the pea is modified when pollen of a differently
coloured variety is used. This conclusion has since been confirmed by
experiments made by the Rev. J. M. Berkeley.[930]
Mr. Laxton of Stamford, whilst making experiments on peas for the
express purpose of ascertaining the influence of foreign pollen on the
mother-plant, has recently[931] observed an important additional fact.
He fertilised the Tall Sugar pea, which bears very thin green pods,
becoming {398} brownish-white when dry, with pollen of the
Purple-podded pea, which, as its name expresses, has dark-purple pods
with very thick skin, becoming pale reddish-purple when dry. Mr. Laxton
has cultivated the tall sugar-pea during twenty years, and has never
seen or heard of it producing a purple pod; nevertheless, a flower
fertilised by pollen from the purple-pod yielded a pod clouded with
purplish-red, which Mr. Laxton kindly gave to me. A space of about two
inches in length towards the extremity of the pod, and a smaller space
near the stalk, were thus coloured. On comparing the colour with that
of the purple-pod, both pods having been first dried and then soaked in
water, it was found to be identically the same; and in both the colour
was confined to the cells lying immediately beneath the outer skin of
the pod. The valves of the crossed pod were also decidedly thicker and
stronger than those of the pods of the mother-plant, but this may have
been an accidental circumstance, for I know not how far their thickness
in the Tall Sugar-pea is a variable character.
The peas of the Tall Sugar-pea, when dry, are pale greenish-brown,
thickly covered with dots of dark purple so minute as to be visible
only through a lens, and Mr. Laxton has never seen or heard of this
variety producing a purple pea; but in the crossed pod one of the peas
was of a uniform beautiful violet-purple tint, and a second was
irregularly clouded with pale purple. The colour lies in the outer of
the two coats which surround the pea. As the peas of the purple-podded
variety when dry are of a pale greenish-buff, it would at first appear
that this remarkable change of colour in the peas in the crossed pod
could not have been caused by the direct action of the pollen of the
purple-pod: but when we bear in mind that this latter variety has
purple flowers, purple marks on its stipules, and purple pods; and that
the Tall sugar-pea likewise has purple flowers and stipules, and
microscopically minute purple dots on the peas, we can hardly doubt
that the tendency to the production of purple in both parents has in
combination modified the colour of the peas in the crossed pod. After
having examined these specimens, I crossed the same two varieties, and
the peas in one pod, but not the pods themselves, were clouded and
tinted with purplish-red in a much more conspicuous manner than the
peas in the uncrossed pods produced at the same time by the same
plants. I may notice as a caution that Mr. Laxton sent me various other
crossed peas slightly, or even greatly, modified in colour; but the
change in these cases was due, as had been suspected by Mr. Laxton, to
the altered colour of the cotyledons, seen through the transparent
coats of the peas; and as the cotyledons are parts of the embryo, these
cases are not in any way remarkable.
Turning now to the genus Matthiola. The pollen of one kind of stock
sometimes affects the colour of the seeds of another kind, used as the
mother-plant. I give the following case the more readily, as Gaertner
doubted similar statements with respect to the stock previously made by
other observers. A well-known horticulturist, Major Trevor Clarke,
informs me[932] that the seeds of the large red-flowered _biennial_
stock {399} (_Matthiola annua_; _Cocardeau_ of the French) are light
brown, and those of the purple branching Queen stock (_M. incana_) are
violet-black; and he found that, when flowers of the red stock were
fertilised by pollen from the purple stock, they yielded about fifty
per cent. of _black_ seeds. He sent me four pods from a red-flowered
plant, two of which had been fertilised by their own pollen, and they
included pale brown seed; and two which had been crossed by pollen from
the purple kind, and they included seeds all deeply tinged with black.
These latter seeds yielded purple-flowered plants like their father;
whilst the pale brown seeds yielded normal red-flowered plants; and
Major Clarke, by sowing similar seeds, has observed on a greater scale
the same result. The evidence in this case of the direct action of the
pollen of one species on the colour of the seeds of another species
appears to me conclusive.
In the foregoing cases, with the exception of that of the purple-podded
pea, the coats of the seeds alone have been affected in colour. We shall
now see that the ovarium itself, whether forming a large fleshy fruit or a
mere thin envelope, may be modified by foreign pollen, in colour, flavour,
texture, size, and shape.
The most remarkable instance, because carefully recorded by highly
competent authorities, is one of which I have seen an account in a
letter written, in 1867, by M. Naudin to Dr. Hooker. M. Naudin states
that he has seen fruit growing on _Chamaerops humilis_, which had been
fertilised by M. Denis with pollen from the Phoenix or date-palm. The
fruit or drupe thus produced was twice as large as, and more elongated
than, that proper to the Chamaerops; so that it was intermediate in
these respects, as well as in texture, between the fruit of the two
parents. These hybridised seeds germinated, and produced young plants
likewise intermediate in character. This case is the more remarkable as
the Chamaerops and Phoenix belong not only to distinct genera, but in
the estimation of some botanists to distinct sections of the family.
Gallesio[933] fertilised the flowers of an orange with pollen from the
lemon; and one fruit thus produced bore a longitudinal stripe of peel
having the colour, flavour, and other characters of the lemon. Mr.
Anderson[934] fertilised a green-fleshed melon with pollen from a
scarlet-fleshed kind; in two of the fruits "a sensible change was
perceptible; and four other fruits were somewhat altered both
internally and externally." The seeds of the two first-mentioned fruits
produced plants partaking of the good properties of both parents. In
the United States, where Cucurbitaceae are largely cultivated, it is the
popular belief[935] that the fruit is thus directly affected by foreign
pollen; and I have received a similar statement with respect to {400}
the cucumber in England. It is known that grapes have been thus
affected in colour, size, and shape: in France a pale-coloured grape
had its juice tinted by the pollen of the dark-coloured Teinturier; in
Germany a variety bore berries which were affected by the pollen of two
adjoining kinds; some of the berries being only partially affected or
mottled.[936] As long ago as 1751[937] it was observed that, when
differently coloured varieties of maize grow near each other, they
mutually affect each other's seeds, and this is now a popular belief in
the United States. Dr. Savi[938] tried the experiment with care: he
sowed yellow and black-seeded maize together, and on the same ear some
of the seeds were yellow, some black, and some mottled,[939] the
differently coloured seeds being arranged in rows or irregularly. Mr.
Sabine states[940] that he has seen the form of the nearly globular
seed-capsule of _Amaryllis vittata_ altered by the application of the
pollen of another species, of which the capsule has gibbous angles. Mr.
J. Anderson Henry[941] crossed _Rhododendron Dalhousiae_ with the pollen
of _R. Nuttallii_, which is one of the largest-flowered and noblest
species of the genus. The largest pod produced by the former species,
when fertilised with its own pollen, measured 1-2/8 inch in length and
11/2 in girth; whilst three of the pods which had been fertilised by
pollen of _R. Nuttallii_ measured 1-5/8 inch in length and no less than
2 inches in girth. Here we see the effect of foreign pollen apparently
confined to increasing the size of the ovarium; but we must be cautious
in assuming, as the following case shows, that in this instance size
has been directly transferred from the male parent to the capsule of
the female plant. Mr. Henry fertilised _Arabis blepharophylla_ with
pollen of _A. Soyeri_, and the pods thus produced, of which he was so
kind as to send me detailed measurements and sketches, were much larger
in all their dimensions than those naturally produced by either the
male or female parent-species. In a future chapter we shall see {401}
that the organs of vegetation in hybrid plants, independently of the
character of either parent, are sometimes developed to a monstrous
size; and the increased size of the pods in the foregoing cases may be
an analogous fact.
No case of the direct action of the pollen of one variety on another is
better authenticated or more remarkable than that of the common apple.
The fruit here consists of the lower part of the calyx and of the upper
part of the flower-peduncle[942] in a metamorphosed condition, so that
the effect of the foreign pollen has extended even beyond the limits of
the ovarium. Cases of apples thus affected were recorded by Bradley in
the early part of the last century; and other cases are given in old
volumes of the Philosophical Transactions;[943] in one of these a
Russeting apple and an adjoining kind mutually affected each other's
fruit; and in another case a smooth apple affected a rough-coated kind.
Another instance has been given[944] of two very different apple-trees
growing close to each other, which bore fruit resembling each other,
but only on the adjoining branches. It is, however, almost superfluous
to adduce these or other cases, after that of the St. Valery apple,
which, from the abortion of the stamens, does not produce pollen, but,
being annually fertilised by the girls of the neighbourhood with pollen
of many kinds, bears fruit, "differing from each other in size,
flavour, and colour, but resembling in character the hermaphrodite
kinds by which they have been fertilised."[945]
I have now shown, on the authority of several excellent observers, in the
case of plants belonging to widely different orders, that the pollen of one
species or variety, when applied to a distinct form, occasionally causes
the coats of the seeds and the ovarium or fruit, including even in one
instance the calyx and upper part of the peduncle of the mother-plant, to
become modified. Sometimes the whole of the ovarium or all the seeds are
thus affected; sometimes only a certain number of the seeds, as in the case
of the pea, or only a part of the ovarium, as with the striped orange,
mottled grapes and maize, are thus affected. It must not be supposed that
any direct or immediate effect invariably follows the use of foreign
pollen: this is far from being the case; nor is it known on what conditions
the result depends. Mr. Knight[946] expressly states that he has never seen
{402} the fruit thus affected, though he has crossed thousands of apple and
other fruit-trees. There is not the least reason to believe that a branch
which has borne seed or fruit directly modified by foreign pollen is itself
affected, so as subsequently to produce modified buds: such an occurrence,
from the temporary connection of the flower with the stem, would be hardly
possible. Hence but very few, if any, of the cases of sudden modifications
in the fruit of trees, given in the early part of this chapter, can be
accounted for by the action of foreign pollen; for such modified fruits
have commonly been afterwards propagated by budding or grafting. It is also
obvious that changes of colour in the flower which necessarily supervene
long before it is ready for fertilisation, and changes in the shape or
colour of the leaves, can have no relation to the action of foreign pollen:
all such cases must be attributed to simple bud-variation.
The proofs of the action of foreign pollen on the mother-plant have been
given in considerable detail, because this action, as we shall see in a
future chapter, is of the highest theoretical importance, and because it is
in itself a remarkable and apparently anomalous circumstance. That it is
remarkable under a physiological point of view is clear, for the male
element not only affects, in accordance with its proper function, the germ,
but the surrounding tissues of the mother-plant. That the action is
anomalous in appearance is true, but hardly so in reality, for apparently
it plays the same part in the ordinary fertilisation of many flowers.
Gaertner has shown,[947] by gradually increasing the number of pollen-grains
until he succeeded in fertilising a Malva, that many grains are expended in
the development, or, as he expresses it, in the satiation, of the pistil
and ovarium. Again, when one plant is fertilised by a widely distinct
species, it often happens that the ovarium is fully and quickly developed
without any seeds being formed, or the coats of the seeds are developed
without an embryo being formed within. Dr. Hildebrand also has lately shown
in a valuable paper[948] that, with several Orchideae, the action of the
plant's own {403} pollen is necessary for the development of the ovarium,
and that this development takes place not only long before the pollen-tubes
have reached the ovules, but even before the placentae and ovules have been
formed; so that with these orchids the pollen apparently acts directly on
the ovarium. On the other hand, we must not overrate the efficacy of pollen
in this respect; for in the case of hybridised plants it might be argued
that an embryo had been formed and had affected the surrounding tissues of
the mother-plant before it perished at a very early age. Again, it is well
known that with many plants the ovarium may be fully developed, though
pollen be wholly excluded. And lastly, Mr. Smith, the late Curator at Kew
(as I hear through Dr. Hooker), observed the singular fact with an orchid,
the _Bonatea speciosa_, the development of the ovarium could be effected by
mechanical irritation of the stigma. Nevertheless, from the number of the
pollen-grains expended "in the satiation of the ovarium and pistil,"--from
the generality of the formation of the ovarium and seed-coats in sterile
hybridised plants,--and from Dr. Hildebrand's observations on orchids, we
may admit that in most cases the swelling of the ovarium, and the formation
of the seed-coats, are at least aided, if not wholly caused, by the direct
action of the pollen, independently of the intervention of the fertilised
germ. Therefore, in the previously-given cases we have only to add to our
belief in the power of the plant's own pollen on the development of the
ovarium and seed-coats, its further power, when applied to a distinct
species or variety, of influencing the shape, size, colour, texture, &c.,
of these same parts.
* * * * *
Turning now to the animal kingdom. If we could imagine the same flower to
yield seeds during successive years, then it would not be very surprising
that a flower of which the ovarium had been modified by foreign pollen
should next year produce, when self-fertilised, offspring modified by the
previous male influence. Closely analogous cases have actually occurred
with animals. In the case often quoted from Lord Morton,[949] a nearly
purely-bred, Arabian, chesnut mare bore a hybrid to a quagga; she was
subsequently sent to Sir Gore Ouseley, and produced {404} two colts by a
black Arabian horse. These colts were partially dun-coloured, and were
striped on the legs more plainly than the real hybrid, or even than the
quagga. One of the two colts had its neck and some other parts of its body
plainly marked with stripes. Stripes on the body, not to mention those on
the legs, and the dun-colour, are extremely rare,--I speak after having
long attended to the subject,--with horses of all kinds in Europe, and are
unknown in the case of Arabians. But what makes the case still more
striking is that the hair of the mane in these colts resembled that of the
quagga, being short, stiff, and upright. Hence there can be no doubt that
the quagga affected the character of the offspring subsequently begot by
the black Arabian horse. With respect to the varieties of our domesticated
animals, many similar and well-authenticated facts have been
published,[950] and others have been communicated to me, plainly showing
the influence of the first male on the progeny subsequently borne by the
mother to other males. It will suffice to give a single instance, recorded
in the 'Philosophical Transactions,' in a paper following that by Lord
Morton: Mr. Giles put a sow of Lord Western's black and white Essex breed
to a wild boar of a deep chesnut colour; and the "pigs produced partook in
appearance of both boar and sow, but in some the chesnut colour of the boar
strongly prevailed." After the boar had long been dead, the sow was put to
a boar of her own black and white breed,--a kind which is well known to
breed very true and never to show any chesnut colour,--yet from this union
the sow produced some young pigs which were plainly marked with the same
chesnut tint as in the first litter. Similar cases have so frequently
occurred, that careful breeders avoid putting a choice female to an
inferior male on account of the injury to her subsequent progeny which may
be expected to follow.
{405}
Some physiologists have attempted to account for these remarkable results
from a first impregnation by the close attachment and freely
intercommunicating blood-vessels between the modified embryo and the
mother. But it is a most improbable hypothesis that the mere blood of one
individual should affect the reproductive organs of another individual in
such a manner as to modify the subsequent offspring. The analogy from the
direct action of foreign pollen on the ovarium and seed-coats of the
mother-plant strongly supports the belief that the male element acts
directly on the reproductive organs of the female, wonderful as is this
action, and not through the intervention of the crossed embryo. With birds
there is no such close connection between the embryo and mother as in the
case of mammals: yet a careful observer, Dr. Chapuis, states[951] that with
pigeons the influence of a first male sometimes makes itself perceived in
the succeeding broods; but this statement, before it can be fully trusted,
requires confirmation.
* * * * *
_Conclusion and Summary of the Chapter._--The facts given in the latter
half of this chapter are well worthy of consideration, as they show us in
how many extraordinary modes one organic form may lead to the modification
of another, and often without the intervention of seminal reproduction.
There is ample evidence, as we have just seen, that the male element may
either directly affect the structure of the female, or in the case of
animals lead to the modification of her offspring. There is a considerable
but insufficient body of evidence showing that the tissues of two plants
may unite and form a bud having a blended character; or again, that buds
inserted into a stock may affect all the buds subsequently produced by this
stock. Two embryos, differing from each other and contained in the same
seed, may cohere and form a single plant. Offspring from a cross between
two species or varieties may in the first or in a succeeding generation
revert in various degrees by bud-variation to their parent-forms; and this
reversion or segregation of character may affect the whole flower, fruit,
or leaf-bud, or only the half or smaller segment, or a single organ. In
some cases this segregation of character apparently depends on some {406}
incapacity of union rather than on reversion, for the flowers or fruit
which are first produced display by segments the characters of both
parents. In the _Cytisus adami_ and the Bizzarria orange, whatever their
origin may have been, the two parent species occur blended together under
the form of a sterile hybrid, or reappear with their characters perfect and
their reproductive organs effective; and these trees, retaining the same
sportive character, can be propagated by buds. These various facts ought to
be well considered by any one who wishes to embrace under a single point of
view the various modes of reproduction by gemmation, division, and sexual
union, the reparation of lost parts, variation, inheritance, reversion, and
other such phenomena. In a chapter towards the close of the following
volume I shall attempt to connect these facts together by a provisional
hypothesis.
In the early half of this chapter I have given a long list of plants in
which through bud-variation, that is, independently of reproduction by
seed, the fruit has suddenly become modified in size, colour, flavour,
hairiness, shape, and time of maturity; flowers have similarly changed in
shape, colour, and doubleness, and greatly in the character of the calyx;
young branches or shoots have changed in colour, in bearing spines, and in
habit of growth, as in climbing and weeping; leaves have changed in colour,
variegation, shape, period of unfolding, and in their arrangement on the
axis. Buds of all kinds, whether produced on ordinary branches or on
subterranean stems, whether simple or, as in tubers and bulbs, much
modified and supplied with a stock of nutriment, are all liable to sudden
variations of the same general nature.
In the list, many of the cases are certainly due to reversion to characters
not acquired from a cross, but which were formerly present, and have been
lost for a longer or shorter period of time;--as when a bud on a variegated
plant produces plain leaves, or when variously-coloured flowers on the
Chrysanthemum revert to the aboriginal yellow tint. Many other cases
included in the list are probably due to the plants being of crossed
parentage, and to the buds reverting to one of the two parent-forms. In
illustration of the origin of _Cytisus adami_, several cases were given of
partial or complete reversion, both {407} with hybrid and mongrel plants;
hence we may suspect that the strong tendency in the Chrysanthemum, for
instance, to produce by bud-variation differently-coloured flowers, results
from the varieties formerly having been intentionally or accidentally
crossed; and that their descendants at the present day still occasionally
revert by buds to the colours of the more persistent parent-varieties. This
is almost certainly the case with Rollisson's Unique Pelargonium; and so it
may be to a large extent with the bud-varieties of the Dahlia and with the
"broken colours" of Tulips.
Many cases of bud-variation, however, cannot be attributed to reversion,
but to spontaneous variability, such as so commonly occurs with cultivated
plants when raised from seed. As a single variety of the Chrysanthemum has
produced by buds six other varieties, and as one variety of the gooseberry
has borne at the same time four distinct varieties of fruit, it is scarcely
possible to believe that all these variations are reversions to former
parents. We can hardly believe, as remarked in a previous chapter, that all
the many peaches which have yielded nectarine-buds are of crossed
parentage. Lastly, in such cases as that of the moss-rose with its peculiar
calyx, and of the rose which bears opposite leaves, in that of the
Imatophyllum, &c., there is no known natural species or seedling variety,
from which the characters in question could have been derived by crossing.
We must attribute all such cases to actual variability in the buds. The
varieties which have thus arisen cannot be distinguished by any external
character from seedlings; this is notoriously the case with the varieties
of the Rose, Azalea, and many other plants. It deserves notice that all the
plants which have yielded bud-variations have likewise varied greatly by
seed.
These plants belong to so many orders that we may infer that almost every
plant would be liable to bud-variation if placed under the proper exciting
conditions. These conditions, as far as we can judge, mainly depend on
long-continued and high cultivation; for almost all the plants in the
foregoing lists are perennials, and have been largely propagated in many
soils and under different climates, by cuttings, offsets, bulbs, tubers,
and especially by budding or grafting. The instances of annuals varying by
buds, or producing on the same plant {408} differently coloured flowers,
are comparatively rare: Hopkirk[952] has seen this with _Convolvulus
tricolor_; and it is not rare with the Balsam and annual Delphinium.
According to Sir R. Schomburgk, plants from the warmer temperate regions,
when cultivated under the hot climate of St. Domingo, are eminently liable
to bud-variation; but change of climate is by no means a necessary
contingent, as we see with the gooseberry, currant, and some others. Plants
living under their natural conditions are very rarely subject to
bud-variation: variegated and coloured leaves have, however, been
occasionally observed; and I have given an instance of the variation of
buds on an ash-tree; but it is doubtful whether any tree planted in
ornamental grounds can be considered as living under strictly natural
conditions. Gaertner has seen white and dark-red flowers produced from the
same root of the wild _Achillea millefolium_; and Prof. Caspary has seen
_Viola lutea_, in a completely wild condition, bearing flowers of different
colours and sizes.[953]
As wild plants are so rarely liable to bud-variation, whilst highly
cultivated plants long propagated by artificial means have yielded by this
form of reproduction many varieties, we are led through a series such as
the following,--namely, all the eyes in the same tuber of the potato
varying in the same manner,--all the fruit on a purple plum-tree suddenly
becoming yellow,--all the fruit on a double-flowered almond suddenly
becoming peach-like,--all the buds on grafted trees being in some very
slight degree affected by the stock on which they have been worked,--all
the flowers on a transplanted heartsease changing for a time in colour,
size, and shape,--we are led through such facts to look at every case of
bud-variation as the direct result of the particular conditions of life to
which the plant has been exposed. But if we turn to the other end of the
series, namely, to such cases as that of a peach-tree which, after having
been cultivated by tens of thousands during many years in many countries,
and after having annually produced thousands of buds, all of which have
apparently been exposed to precisely the same conditions, yet at last
suddenly produces a single bud with its whole character greatly
transformed, we are driven to an opposite {409} conclusion. In such cases
as the latter it would appear that the transformation stands in no _direct_
relation to the conditions of life.
We have seen that varieties produced from seeds and from buds resemble each
other so closely in general appearance, that they cannot possibly be
distinguished. Just as certain species and groups of species, when
propagated by seed, are more variable than other species or genera, so it
is in the case of certain bud-varieties. Thus the Queen of England
Chrysanthemum has produced by this latter process no less than six, and
Rollisson's Unique Pelargonium four distinct varieties; moss-roses have
also produced several other moss-roses. The Rosaceae have varied by buds
more than any other group of plants; but this may be in large part due to
so many members having been long cultivated; but within this one group, the
peach has often varied by buds, whilst the apple and pear, both grafted
trees extensively cultivated, have afforded, as far as I can ascertain,
extremely few instances of bud-variation.
The law of analogous variation holds good with varieties produced by buds,
as with those produced from seed: more than one kind of rose has sported
into a moss-rose; more than one kind of camellia has assumed an hexagonal
form; and at least seven or eight varieties of the peach have produced
nectarines.
The laws of inheritance seem to be nearly the same with seminal and
bud-varieties. We know how commonly reversion comes into play with both,
and it may affect the whole, or only segments, of a leaf, flower, or fruit.
When the tendency to reversion affects many buds on the same tree, it
becomes covered with different kinds of leaves, flowers, or fruit; but
there is reason to believe that such fluctuating varieties have generally
arisen from seed. It is well known that, out of a number of seedling
varieties, some transmit their character much more truly by seed than
others; so with bud-varieties some retain their character by successive
buds more truly than others; of which instances have been given with two
kinds of variegated Euonymus and with certain kinds of tulips.
Notwithstanding the sudden production of bud-varieties, the characters thus
acquired are sometimes capable of transmission by seminal reproduction: Mr.
Rivers has found that moss-roses generally {410} reproduce themselves by
seed; and the mossy character has been transferred by crossing, from one
species of rose to another. The Boston nectarine, which appeared as a
bud-variation, produced by seed a closely allied nectarine. We have however
seen, on the authority of Mr. Salter, that seed taken from a branch with
leaves variegated through bud-variation, transmits this character very
feebly; whilst many plants, which became variegated as seedlings, transmit
variegation to a large proportion of their progeny.
Although I have been able to collect a good many cases of bud-variation, as
shown in the previous lists, and might probably, by searching foreign
horticultural works, have collected more cases, yet their total number is
as nothing in comparison with that of seminal varieties. With seedlings
raised from the more variable cultivated plants, the variations are almost
infinitely numerous, but their differences are generally slight: only at
long intervals of time a strongly marked modification appears. On the other
hand, it is a singular and inexplicable fact that, when plants vary by
buds, the variations, though they occur with comparative rarity, are often,
or even generally, strongly pronounced. It struck me that this might
perhaps be a delusion, and that slight changes often occurred in buds, but
from being of no value were overlooked or not recorded. Accordingly I
applied to two great authorities on this subject, namely, to Mr. Rivers
with respect to fruit-trees, and to Mr. Salter with respect to flowers. Mr.
Rivers is doubtful, but does not remember having noticed very slight
variations in fruit-buds. Mr. Salter informs me that with flowers such do
occur, but, if propagated, they generally lose their new character in the
following year; yet he concurs with me that bud-variations usually at once
assume a decided and permanent character. We can hardly doubt that this is
the rule, when we reflect on such cases as that of the peach, which has
been so carefully observed and of which such trifling seminal varieties
have been propagated, yet this tree has repeatedly produced by
bud-variation nectarines, and only twice (as far as I can learn) any other
variety, namely, the Early and Late Grosse Mignonne peaches; and these
differ from the parent-tree in hardly any character except the period of
maturity. {411}
To my surprise I hear from Mr. Salter that he brings the great principle of
selection to bear on variegated plants propagated by buds, and has thus
greatly improved and fixed several varieties. He informs me that at first a
branch often produces variegated leaves on one side alone, and that the
leaves are marked only with an irregular edging or with a few lines of
white and yellow. To improve and fix such varieties, he finds it necessary
to encourage the buds at the bases of the most distinctly marked leaves,
and to propagate from them alone. By following with perseverance this plan
during three or four successive seasons, a distinct and fixed variety can
generally be secured.
Finally, the facts given in this chapter prove in how close and remarkable
a manner the germ of a fertilised seed and the small cellular mass forming
a bud resemble each other in function,--in their powers of inheritance with
occasional reversion,--and in their capacity for variation of the same
general nature, in obedience to the same laws. This resemblance, or rather
identity, is rendered far more striking if the facts can be trusted which
apparently render it probable that the cellular tissue of one species or
variety, when budded or grafted on another, may give rise to a bud having
an intermediate character. In this chapter we clearly see that variability
is not necessarily contingent on sexual generation, though much more
frequently its concomitant than on bud-reproduction. We see that
bud-variability is not solely dependent on reversion or atavism to
long-lost characters, or to those formerly acquired from a cross, but that
it is often spontaneous. But when we ask ourselves what is the cause of any
particular bud-variation, we are lost in doubt, being driven in some cases
to look to the direct action of the external conditions of life as
sufficient, and in other cases to feel a profound conviction that these
have played a quite subordinate part, of not more importance than the
nature of the spark which ignites a mass of combustible matter.
END OF VOL. I.
LONDON:
PRINTED BY W. CLOWES AND SONS, STAMFORD STREET,
AND CHARING CROSS.
* * * * *
NOTES
[1] To any one who has attentively read my 'Origin of Species' this
Introduction will be superfluous. As I stated that work that I should soon
publish the facts on which the conclusions given in it were founded, I here
beg permission to remark that the great delay in publishing this first work
has been caused by continued ill-health.
[2] M. Pouchet has recently ('Plurality of Races,' Eng. Translat., 1864, p.
83, &c.) insisted that variation under domestication throws no light on the
natural modification of species. I cannot perceive the force of his
arguments, or, to speak more accurately, of his assertions to this effect.
[3] Leon Dufour in 'Annales des Scienc. Nat.' (3rd series, Zoolog.), tom.
v. p. 6.
[4] In treating the several subjects included in the present and succeeding
works I have continually been led to ask for information from many
zoologists, botanists, geologists, breeders of animals, and
horticulturists, and I have invariably received from them the most generous
assistance. Without such aid I could have effected little. I have
repeatedly applied for information and specimens to foreigners, and to
British merchants and officers of the Government residing in distant lands,
and, with the rarest exceptions, I have received prompt, open-handed, and
valuable assistance. I cannot express too strongly my obligations to the
many persons who have assisted me, and who, I am convinced, would be
equally willing to assist others in any scientific investigation.
[5] Owen, 'British Fossil Mammals,' p. 123 to 133. Pictet's 'Traite de
Pal.,' 1853, tom. i. p. 202. De Blainville, in his 'Osteographie, Canidae,'
p. 142, has largely discussed the whole subject, and concludes that the
extinct parent of all domesticated dogs came nearest to the wolf in
organization, and to the jackal in habits.
[6] Pallas, I believe, originated this doctrine in 'Act. Acad. St.
Petersburgh,' 1780, Part ii. Ehrenberg has advocated it, as may be seen in
De Blainville's 'Osteographie,' p. 79. It has been carried to an extreme
extent by Col. Hamilton Smith in the 'Naturalist Library,' vol. ix. and x.
Mr. W. C. Martin adopts it in his excellent 'History of the Dog,' 1845; as
does Dr. Morton, as well as Nott and Gliddon, in the United States. Prof.
Low, in his 'Domesticated Animals,' 1845, p. 666, comes to this same
conclusion. No one has argued on this side with more clearness and force
than the late James Wilson, of Edinburgh, in various papers read before the
Highland Agricultural and Wernerian Societies. Isidore Geoffroy Saint
Hilaire ('Hist. Nat. Gen.,' 1860, tom. iii. p. 107), though he believes
that most dogs have descended from the jackal, yet inclines to the belief
that some are descended from the wolf. Prof. Gervais ('Hist. Nat. Mamm.,'
1855, tom. ii. p. 69), referring to the view that all the domestic races
are the modified descendants of a single species, after a long discussion,
says, "Cette opinion est, suivant nous du moins, la moins probable."
[7] Berjeau, 'The Varieties of the Dog; in old Sculptures and Pictures,'
1863. 'Der Hund,' von Dr. F. L. Walther, s. 48, Giessen, 1817: this author
seems carefully to have studied all classical works on the subject. _See_
also 'Volz, Beitraege zur Kultur-geschichte,' Leipzig, 1852, s. 115. 'Youatt
on the Dog,' 1845, p. 6. A very full history is given by De Blainville in
his 'Osteographie, Canidae.'
[8] I have seen drawings of this dog from the tomb of the son of Esar
Haddon, and clay models in the British Museum. Nott and Gliddon, in their
'Types of Mankind,' 1854, p. 393, give a copy of these drawings. This dog
has been called a Thibetan mastiff, but Mr. H. A. Oldfield, who is familiar
with the so-called Thibet mastiff, and has examined the drawings in the
British Museum, informs me that he considers them different.
[9] 'Proc. Zoolog. Soc.,' July 12th, 1831.
[10] 'Sporting in Algeria,' p. 51.
[11] Berjeau gives fac-similes of the Egyptian drawings. Mr. C. L. Martin,
in his 'History of the Dog,' 1845, copies several figures from the Egyptian
monuments, and speaks with much confidence with respect to their identity
with still living dogs. Messrs. Nott and Gliddon ('Types of Mankind,' 1854,
p. 388) give still more numerous figures. Mr. Gliddon asserts that a
curl-tailed greyhound, like that represented on the most ancient monuments,
is common in Borneo; but the Rajah, Sir J. Brooke, informs me that no such
dog exists there.
[12] These, and the following facts on the Danish remains, are taken from
M. Morlot's most interesting memoir in 'Soc. Vaudoise des Sc. Nat.,' tom.
vi., 1860, pp. 281, 299, 320.
[13] 'Die Fauna der Pfahlbauten,' 1861, s. 117, 162.
[14] De Blainville, 'Osteographie, Canidae.'
[15] Sir R. Schomburgk has given me information on this head. _See_ also
'Journal of R. Geograph. Soc.,' vol. xiii., 1843, p. 65.
[16] 'Domestication of Animals:' Ethnological Soc., Dec. 22nd, 1863.
[17] 'Journal of Researches,' &c., 1845, p. 393. With respect to _Canis
antarcticus_, _see_ p. 193. For the case of the antelope, _see_ 'Journal
Royal Geograph. Soc.,' vol. xxiii. p. 94.
[18] The authorities for the foregoing statements are as
follow:--Richardson, in 'Fauna Boreali-Americana,' 1829, pp. 64, 75; Dr.
Kane, 'Arctic Explorations,' 1856, vol. i. pp. 398, 455; Dr. Hayes, 'Arctic
Boat Journey,' 1860, p. 167. Franklin's 'Narrative,' vol. i. p. 269, gives
the case of three whelps of a black wolf being carried away by the Indians.
Parry, Richardson, and others, give accounts of wolves and dogs naturally
crossing in the eastern parts of North America. Seeman, in his 'Voyage of
H.M.S. Herald,' 1853, vol. ii. p. 26, says the wolf is often caught by the
Esqimaux for the purpose of crossing with their dogs, and thus adding to
their size and strength. M. Lamare-Picquot, in 'Bull. de la Soc.
d'Acclimat.,' tom. vii., 1860, p. 148, gives a good account of the
half-bred Esquimaux dogs.
[19] 'Fauna Boreali-Americana,' 1829, pp. 73, 78, 80. Nott and Gliddon,
'Types of Mankind,' p. 383. The naturalist and traveller Bartram is quoted
by Hamilton Smith, in 'Nat. Hist. Lib.,' vol. x. p. 156. A Mexican domestic
dog seems also to resemble a wild dog of the same country; but this may be
the prairie-wolf. Another capable judge, Mr. J. K. Lord ('The Naturalist in
Vancouver Island,' 1866, vol. ii. p. 218), says that the Indian dog of the
Spokans, near the Rocky Mountains, "is beyond all question nothing more
than a tamed Cayote or prairie-wolf," or _Canis latrans_.
[20] I quote this from Mr. R. Hill's excellent account of the Alco or
domestic dog of Mexico, in Gosse's 'Naturalist's Sojourn in Jamaica,' 1851,
p. 329.
[21] 'Naturgeschichte der Saeugethiere von Paraguay,' 1830, s. 151.
[22] Quoted in Humboldt's 'Aspects of Nature' (Eng. transl.), vol. i. p.
108.
[23] Paget's 'Travels in Hungary and Transylvania,' vol. i. p. 501.
Jeitteles, 'Fauna Hungariae Superioris,' 1862, s. 13. _See_ Pliny, 'Hist. of
the World' (Eng. transl.), 8th book, ch. xl., about the Gauls crossing
their dogs. _See_ also 'Hist. Animal.' lib. viii. c. 28. For good evidence
about wolves and dogs naturally crossing near the Pyrenees, _see_ M.
Mauduyt, 'Du Loup et de ses Races,' Poitiers, 1851; also Pallas, in 'Acta
Acad. St. Petersburgh,' 1780, part ii. p. 94.
[24] I give this on excellent authority, namely, Mr. Blyth (under the
signature of Zoophilus), in the 'Indian Sporting Review,' Oct. 1856, p.
134. Mr. Blyth states that he was struck with the resemblance between a
brush-tailed race of pariah-dogs, north-west of Cawnpore, and the Indian
wolf. He gives corroborative evidence with respect to the dogs of the
valley of the Nerbudda.
[25] For numerous and interesting details on the resemblance of dogs and
jackals, _see_ Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' 1860, tom.
iii. p. 101. _See_ also 'Hist. Nat. des Mammiferes,' par Prof. Gervais,
1855, tom. ii. p. 60.
[26] Gueldenstaedt, 'Nov. Comment. Acad. Petrop.,' tom. xx., pro anno 1775,
p. 449.
[27] Quoted by De Blainville in his 'Osteographie, Canidae,' pp. 79, 98.
[28] _See_ Pallas, in 'Act. Acad. St. Petersburgh,' 1780, part ii. p. 91.
For Algeria, _see_ Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii.
p. 177. In both countries it is the male jackal which pairs with female
domestic dogs.
[29] John Barbut's 'Description of the Coast of Guinea in 1746.'
[30] 'Travels in South Africa,' vol. ii. p. 272.
[31] Selwyn, Geology of Victoria; 'Journal of Geolog. Soc.,' vol. xiv.,
1858, p. 536, and vol. xvi., 1860, p. 148; and Prof M^cCoy, in 'Annals and
Mag. of Nat. Hist.' (3rd series), vol. ix., 1862, p. 147. The Dingo differs
from the dogs of the central Polynesian islands. Dieffenbach remarks
('Travels,' vol. ii. p. 45) that the native New Zealand dog also differs
from the Dingo.
[32] 'Proceedings Zoolog. Soc.,' 1833, p. 112. _See_, also, on the taming
of the common wolf, L. Lloyd, 'Scandinavian Adventures,' vol. i. p. 460,
1854. With respect to the jackal, _see_ Prof. Gervais, 'Hist. Nat. Mamm.,'
tom. ii. p. 61. With respect to the aguara of Paraguay, _see_ Rengger's
work.
[33] Roulin, in 'Mem. present. par divers Savans,' tom. vi. p. 341.
[34] Martin, 'History of the Dog,' p. 14.
[35] Quoted by L. Lloyd in 'Field Sports of North of Europe,' vol. i. p.
387.
[36] Quatrefages, 'Soc. d'Acclimat.,' May 11th, 1863, p. 7.
[37] 'Annals and Mag. of Nat. Hist.,' vol. xv., 1845, p. 140.
[38] Azara, 'Voyages dans l'Amer. Merid.,' tom. i. p. 381; his account is
fully confirmed by Rengger. Quatrefages gives an account of a bitch brought
from Jerusalem to France which burrowed a hole and littered in it. _See_
'Discours, Exposition des Races Canines,' 1865, p. 3.
[39] With respect to wolves burrowing holes, _see_ Richardson, Fauna
Boreali-Americana,' p. 64; and Bechstein, 'Naturgesch. Deutschlands,' b. i.
s. 617.
[40] _See_ Poeppig, 'Reise in Chile,' b. i. s. 290; Mr. G. Clarke, as
above; and Rengger, s. 155.
[41] Dogs, 'Nat. Library,' vol. x. p. 121: an endemic South American dog
seems also to have become feral in this island. _See_ Gosse's 'Jamaica,' p.
340.
[42] Low, 'Domesticated Animals,' p. 650.
[43] 'The Naturalist Library,' Dogs, vol. x. pp. 4, 19.
[44] Quoted by Prof. Gervais, 'Hist. Nat. Mamm.,' tom. ii. p. 66.
[45] J. Hunter shows that the long period of seventy-three days given by
Buffon is easily explained by the bitch having received the dog many times
during a period of sixteen days ('Phil. Transact.,' 1787, p. 253). Hunter
found that the gestation of a mongrel from wolf and dog ('Phil. Transact.,'
1759, p. 160) apparently was sixty-three days, for she received the dog
more than once. The period of a mongrel dog and jackal was fifty-nine days.
Fred. Cuvier found the period of gestation of the wolf to be ('Dict. Class.
d'Hist. Nat.,' tom. iv. p. 8) two months and a few days, which agrees with
the dog. Isid. G. St. Hilaire, who has discussed the whole subject, and
from whom I quote Bellingeri, states ('Hist. Nat. Gen.,' tom. iii. p. 112)
that in the Jardin des Plantes the period of the jackal has been found to
be from sixty to sixty-three days, exactly as with the dog.
[46] _See_ Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 112,
on the odour of jackals. Col. Ham. Smith, in 'Nat. Hist. Lib.,' vol. x. p.
289.
[47] Quoted by Quatrefages in 'Bull. Soc. d'Acclimat.,' May 11th, 1863.
[48] 'Journal de la Physiologie,' tom. ii. p. 385.
[49] _See_ Mr. R. Hill's excellent account of this breed in Gosse's
'Jamaica,' p. 338; Rengger's 'Saeugethiere von Paraguay,' s. 153. With
respect to Spitz dogs, _see_ Bechstein's 'Naturgesch. Deutschlands,' 1801,
b. i. s. 638. With respect to Dr. Hodgkin's statement made before Brit.
Assoc., _see_ 'The Zoologist,' vol. iv., for 1845-46, p. 1097.
[50] 'Acta Acad. St. Petersburgh,' 1780, part ii. pp. 84, 100.
[51] M. Broca has shown ('Journal de Physiologie,' tom. ii. p. 353) that
Buffon's experiments have been often misrepresented. Broca has collected
(pp. 390-395) many facts on the fertility of crossed dogs, wolves, and
jackals.
[52] 'De la Longevite Humaine,' par M. Flourens, 1855, p. 143. Mr. Blyth
says ('Indian Sporting Review,' vol. ii. p. 137) that he has seen in India
several hybrids from the pariah-dog and jackal; and between one of these
hybrids and a terrier. The experiments of Hunter on the jackal are well
known. See also Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii, p.
217, who speaks of the hybrid offspring of the jackal as perfectly fertile
for three generations.
[53] On authority of F. Cuvier, quoted in Bronn's 'Geschichte der Natur,'
B. ii. s. 164.
[54] W. C. L. Martin, 'History of the Dog,' 1845, p. 203. Mr. Philip P.
King, after ample opportunities of observation, informs me that the Dingo
and European dogs often cross in Australia.
[55] Rueppel, 'Neue Wirbelthiere von Abyssinien,' 1835-40; 'Mammif.,' s. 39,
pl. xiv. There is a specimen of this fine animal in the British Museum.
[56] Even Pallas admits this: _see_ 'Act. Acad. St. Petersburgh,' 1780, p.
93.
[57] Quoted by I. Geoffroy, 'Hist. Nat. Gen.,' tom. iii. p. 453.
[58] F. Cuvier, in 'Annales du Museum,' tom. xviii. p. 337; Godron, 'De
l'Espece,' tom. i. p. 342; and Col. Ham. Smith, in 'Naturalist's Library,'
vol. ix. p. 101.
[59] Isid. Geoffroy Saint Hilaire, 'Hist. des Anomalies,' 1832, tom. i. p.
660. Gervais, 'Hist. Nat. des Mammiferes,' tom. ii., 1855, p. 66. De
Blainville ('Osteographie, Canidae,' p. 137) has also seen an extra molar on
both sides.
[60] 'Osteographie, Canidae,' p. 137.
[61] Wuerzburger, 'Medecin, Zeitschrift,' 1860, B. i. s. 265.
[62] Mr. Yarell, in 'Proc. Zoolog. Soc.,' Oct. 8th, 1833. Mr. Waterhouse
showed me a skull of one of these dogs, which had only a single molar on
each side and some imperfect incisors.
[63] Quoted in 'The Veterinary,' London, vol. viii. p. 415.
[64] 'Hist Nat. General,' tom. iii. p. 448.
[65] W. Scrope, 'Art of Deer-Stalking,' p. 354.
[66] Quoted by Col. Ham. Smith in 'Naturalist's Library,' vol. x. p. 79.
[67] De Blainville, 'Osteographie, Canidae,' p. 134. F. Cuvier, 'Annales du
Museum,' tom. xviii. p. 342. In regard to mastiffs, see Col. Ham. Smith,
'Nat Lib.,' vol. x. p. 218. For the Thibet mastiff, see Mr. Hodgson in
'Journal of As. Soc. of Bengal,' vol. i., 1832, p. 342.
[68] 'The Dog,' 1845, p. 186. With respect to diseases, Youatt asserts (p.
167) that the Italian greyhound is "strongly subject" to polypi in the
matrix or vagina. The spaniel and pug (p. 182) are most liable to
bronchocele. The liability to distemper (p. 232) is extremely different in
different breeds. On the distemper, _see_ also Col. Hutchinson on 'Dog
Breaking,' 1850, p. 279.
[69] _See_ Youatt on the Dog, p. 15; 'The Veterinary,' London, vol. xi. p.
235.
[70] 'Journal of As. Soc. of Bengal,' vol. iii. p. 19.
[71] 'Travels,' vol. ii. p. 15.
[72] Hodgson, in 'Journal of As. Soc. of Bengal,' vol. i. p. 342.
[73] 'Field Sports of the North of Europe,' vol. ii. p. 165.
[74] 'Hist. Nat. des Mammif., 1855, tom. ii. pp. 66, 67.
[75] 'History of Quadrupeds,' 1793, vol. i. p. 238.
[76] 'Oriental Field Sports,' quoted by Youatt, 'The Dog,' p. 15.
[77] Quoted by Mr. Galton, 'Domestication of Animals,' p. 13.
[78] 'Hist. Nat. Gen.,' tom. iii. p. 450.
[79] Mr. Greenhow on the Canadian Dog, in Loudon's 'Mag. of Nat. Hist.,'
vol. vi., 1833, p. 511.
[80] _See_ Mr. C. O. Groom-Napier on the webbing of the hind feet of
Otter-hounds, in 'Land and Water,' Oct. 13th, 1866, p. 270.
[81] 'Fauna Boreali-Americana,' 1829, p. 62.
[82] 'The Horse in all his Varieties,' &c., 1829, pp. 230, 234.
[83] 'The Dog,' 1845, pp. 31, 35; with respect to King Charles's spaniel,
p. 45; for the setter, p. 90.
[84] In the 'Encyclop. of Rural Sports,' p. 557.
[85] 'The Farrier,' 1828, vol. i. p. 337.
[86] _See_ Col. Hamilton Smith on the antiquity of the Pointer, in
'Naturalist's Library,' vol. x. p. 195.
[87] The Newfoundland dog is believed to have originated from a cross
between the Esquimaux dog and a large French hound. _See_ Dr. Hodgkin,
'Brit. Assoc.,' 1844; Bechstein's 'Naturgesch. Deutschlands,' Band i. s.
574; 'Naturalist's Library,' vol. x. p. 132; also Mr. Jukes' 'Excursion in
and about Newfoundland.'
[88] De Blainville, 'Osteographie, Felis,' p. 65, on the character of _F.
caligulata_; pp. 85, 89, 90, 175, on the other mummied species. He quotes
Ehrenberg on _F. maniculata_ being mummied.
[89] Asiatic Soc. of Calcutta; Curator's Report, Aug. 1856. The passage
from Sir W. Jardine is quoted from this Report. Mr. Blyth, who has
especially attended to the wild and domestic cats of India, has given in
this Report a very interesting discussion on their origin.
[90] 'Fauna Hungariae Sup.,' 1862, s. 12.
[91] Isid. Geoffrey Saint Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 177.
[92] 'Proc. Zoolog. Soc.,' 1863, p. 184.
[93] 'Saeugethiere von Paraguay,' 1830, s. 212.
[94] 'Mem. presentes par divers Savans: Acad. Roy. des Sciences,' tom. vi.
p. 346. Gomara first noticed this fact in 1554.
[95] 'Narrative of Voyages,' vol. ii. p. 180.
[96] J. Crawfurd, 'Descript. Dict. of the Indian Islands,' p. 255. The
Madagascar cat is said to have a twisted tail: _see_ Desmarest, in
'Encyclop. Nat. Mamm.,' 1820, p. 233, for some of the other breeds.
[97] Admiral Lutke's Voyage, vol. iii. p. 308.
[98] 'Zoology of the Voyage of the Beagle, Mammalia,' p. 20. Dieffenbach,
'Travels in New Zealand,' vol. ii. p. 185. Ch. St. John, 'Wild Sports of
the Highlands,' 1846, p. 49.
[99] Quoted by Isid. Geoffroy, 'Hist. Nat. Gen.,' tom. iii. p. 427.
[100] Ruetimeyer, 'Fauna der Pfalbauten,' 1861, s. 122.
[101] _See_ Youatt on the Horse: J. Lawrence on the Horse, 1829: W. C. L.
Martin, 'History of the Horse,' 1845: Col. Ham. Smith, in 'Naturalist's
Library, Horses,' 1841, vol. xii.: Prof. Veith, 'Die Naturgesch.
Haussaeugethiere,' 1856.
[102] Crawfurd, 'Descript. Dict. of Indian Islands,' 1856, p. 153. "There
are many different breeds, every island having at least one peculiar to
it." Thus in Sumatra there are at least two breeds; in Achin and Batubara
one; in Java several breeds; one in Bali, Lomboc, Sumbawa (one of the best
breeds), Tambora, Bima, Gunung-api, Celebes, Sumba, and Philippines. Other
breeds are specified by Zollinger in the 'Journal of the Indian
Archipelago,' vol. v. p. 343, &c.
[103] 'The Horse,' &c., by John Lawrence, 1829, p. 14.
[104] 'The Veterinary,' London, vol. v. p. 543.
[105] Proc. Veterinary Assoc., in 'The Veterinary,' vol. xiii. p. 42.
[106] 'Bulletin de la Soc. Geolog.,' tom. xxii., 1866, p. 22.
[107] Mr. Percival, of the Enniskillen Dragoons, in 'The Veterinary,' vol.
i. p. 224: _see_ Azara, 'Des Quadrupedes du Paraguay,' tom. ii. p. 313. The
French translator of Azara refers to other cases mentioned by Huzard as
occurring in Spain.
[108] Godron, 'De l'Espece,' tom i. p. 378.
[109] 'Ueber die Eigenschaften,' &c., 1828, s. 10.
[110] 'Domesticated Animals of the British Islands,' pp. 527, 532. In all
the veterinary treatises and papers which I have read, the writers insist
in the strongest terms on the inheritance by the horse of all good and bad
tendencies and qualities. Perhaps the principle of inheritance is not
really stronger in the horse than in any other animal; but, from its value,
the tendency has been more carefully observed.
[111] Andrew Knight crossed breeds so different in size as a dray-horse and
Norwegian pony: see A. Walker on 'Intermarriage,' 1838, p. 205.
[112] 'Naturalist's Library,' Horses, vol. xii. p. 208.
[113] Gervais, 'Hist Nat. Mamm.,' tom. ii. p. 143. Owen, 'British Fossil
Mammals,' p. 383.
[114] 'Kenntniss der fossilen Pferde,' 1863, s. 131.
[115] Mr. W. C. L. Martin ('The Horse,' 1845, p. 34), in arguing against
the belief that the wild Eastern horses are merely feral, has remarked on
the improbability of man in ancient times having extirpated a species in a
region where it can now exist in numbers.
[116] 'Transact. Maryland Academy,' vol. i. part i. p. 28.
[117] Mr. Mackinnon on 'The Falkland Islands,' p. 25. The average height of
the Falkland horses is said to be 14 hands 2 inches. _See_ also my 'Journal
of Researches.'
[118] Pallas, 'Act. Acad. St. Petersburgh,' 1777, part ii. p. 265. With
respect to the tarpans scraping away the snow, _see_ Col. Hamilton Smith in
'Nat. Lib.,' vol. xii. p. 165.
[119] Franklin's 'Narrative,' vol. i. p. 87; note by Sir J. Richardson.
[120] Mr. J. H. Moor, 'Notices of the Indian Archipelago:' Singapore, 1837,
p. 189. A pony from Java was sent ('Athenaeum,' 1842, p. 718) to the Queen
only 28 inches in height. For the Loo Choo Islands, _see_ Beechey's
'Voyage,' 4th edit., vol. i. p. 499.
[121] J. Crawford, 'History of the Horse;' 'Journal of Royal United Service
Institution,' vol. iv.
[122] 'Essays on Natural History,' 2nd series, p. 161.
[123] 'Quadrupedes du Paraguay,' tom. ii. p. 333.
[124] Prof. Low, 'Domesticated Animals,' p. 546. With respect to the writer
in India, _see_ 'India Sporting Review,' vol. ii. p. 181. As Lawrence has
remarked ('The Horse,' p. 9), "perhaps no instance has ever occurred of a
three-part bred horse (_i.e._ a horse, one of whose grand-parents was of
impure blood) saving his distance in running two miles with thoroughbred
racers." Some few instances are on record of seven-eighths racers having
been successful.
[125] Prof. Gervais (in his 'Hist. Nat. Mamm.,' tom. ii. p. 144) has
collected many facts on this head. For instance, Solomon (Kings, b. i. ch.
x. v. 28) bought horses in Egypt at a high price.
[126] 'The Field,' July 13th, 1861, p. 42.
[127] E. Vernon Harcourt, 'Sporting in Algeria,' p. 26.
[128] I state this from my own observations made during several years on
the colours of horses. I have seen cream-coloured, light-dun and mouse-dun
horses dappled, which I mention because it has been stated (Martin,
'History of the Horse,' p. 134) that duns are never dappled. Martin (p.
205) refers to dappled asses. In 'The Farrier' (London, 1828, pp. 453, 455)
there are some good remarks on the dappling of horses; and likewise in Col.
Hamilton Smith on 'The Horse.'
[129] Some details are given in 'The Farrier,' 1828, pp. 452, 455. One of
the least ponies I ever saw, of the colour of a mouse, had a conspicuous
spinal stripe. A small Indian chesnut pony had the same stripe, as had a
remarkably heavy chesnut cart-horse. Race-horses often have the spinal
stripe.
[130] I have received information, through the kindness of the
Consul-General, Mr. J. R. Crowe, from Prof. Boeck, Rasck, and Esmarck, on
the colours of the Norwegian ponies. _See_, also, 'The Field,' 1861, p.
431.
[131] Col. Ham. Smith, 'Nat. Lib.,' vol. xii. p. 275.
[132] Mr. G. Clark, in 'Annal and Mag. of Nat. History,' 2nd series, vol.
ii., 1848, p. 363. Mr. Wallace informs me that he saw in Java a dun and
clay-coloured horse with spinal and leg stripes.
[133] _See_, also, on this point, 'The Field,' July 27th, 1861, p. 91.
[134] 'The Field,' 1861, pp. 431, 493, 545.
[135] 'Ueber die Eigenschaften,' &c, 1828, s. 13, 14.
[136] 'Naturalist's Library,' vol. xii. (1841), pp. 109, 156 to 163, 280,
281. Cream-colour, passing into Isabella (_i.e._ the colour of the dirty
linen of Queen Isabella), seems to have been common in ancient times. _See_
also Pallas's account of the wild horses of the East, who speaks of dun and
brown as the prevalent colours.
[137] Azara, 'Quadrupedes du Paraguay,' tom. ii. p. 307; for the colour of
mules, _see_ p. 350. In North America, Catlin (vol. ii. p. 57) describes
the wild horses, believed to have descended from the Spanish horses of
Mexico, as of all colours, black, grey, roan, and roan pied with sorrel. F.
Michaux ('Travels in North America,' Eng. translat., p. 235) describes two
wild horses from Mexico as roan. In the Falkland Islands, where the horse
has been feral only between 60 and 70 years, I was told that roans and
iron-greys were the prevalent colours. These several facts show that horses
do not generally revert to any uniform colour.
[138] Dr. Sclater, in 'Proc. Zoolog. Soc.,' 1862, p. 164.
[139] W. C. Martin, 'History of the Horse,' 1845, p. 207.
[140] Col. Sykes' Cat. of Mammalia, 'Proc. Zoolog. Soc.,' July 12th, 1831.
Williamson, 'Oriental Field Sports,' vol. ii., quoted by Martin, p. 206.
[141] Blyth, in 'Charlesworth's Mag. of Nat. Hist.,' vol. iv., 1840, p. 83.
I have also been assured by a breeder that this is the case.
[142] One case is given by Martin, 'The Horse,' p. 205.
[143] 'Journal As. Soc. of Bengal,' vol. xxviii. 1860, p. 231. Martin on
the Horse, p. 205.
[144] Hermann von Nathusius, 'Die Racen des Schweines,' Berlin, 1860; and
'Vorstudien fur Geschichte,' &c., 'Schweineschaedel,' Berlin, 1864.
Ruetimeyer, 'Die Fauna der Pfahlbauten,' Basel, 1861.
[145] Nathusius, 'Die Racen des Schweines,' Berlin, 1860. An excellent
appendix is given with references to published and trustworthy drawings of
the breeds of each country.
[146] For Europe, _see_ Bechstein, 'Naturgesch. Deutschlands,' 1801, b. i.,
s. 505. Several accounts have been published on the fertility of the
offspring from wild and tame swine. _See_ Burdach's 'Physiology,' and
Godron, 'De l'Espece,' tom. i. p. 370. For Africa, 'Bull. de la Soc.
d'Acclimat.,' tom. iv. p. 389. For India, _see_ Nathusius,
'Schweineschaedel,' s. 148.
[147] Sir W. Elliot, Catalogue of Mammalia, 'Madras Journal of Lit. and
Science,' vol. x. p. 219.
[148] 'Pfahlbauten,' s. 163 et passim.
[149] _See_ Ruetimeyer's Neue Beitrage, ... Torfschweine, Verh. Naturfor.
Gesell. in Basel, iv. i., 1865, s. 139.
[150] Stan. Julien, quoted by De Blainville, 'Osteographie,' p. 163.
[151] Richardson, 'Pigs, their Origin,' &c., p. 26.
[152] 'Die Racen des Schweines,' s. 47, 64.
[153] 'Proc. Zoolog. Soc.,' 1861, p. 263.
[154] Sclater, in 'Proc. Zoolog. Soc.,' Feb. 26th, 1861.
[155] 'Proc. Zoolog. Soc.,' 1862, p. 13.
[156] 'Journal of Voyages and Travels from 1821 to 1829,' vol. i. p. 300.
[157] Rev. G. Low, 'Fauna Orcadensis,' p. 10. _See_ also Dr. Hibbert's
account of the pig of the Shetland Islands.
[158] 'Die Racen des Schweines,' s. 70.
[159] These woodcuts are copied from engravings given in Mr. S. Sidney's
excellent edition of 'The Pig,' by Youatt, 1860. _See_ pp. 1, 16, 19.
[160] 'Schweineschaedel,' s. 74, 135.
[161] Nathusius, 'Die Racen des Schweines,' s. 71.
[162] 'Die Racen des Schweines,' s. 47. 'Schweineschaedel,' s. 104. Compare,
also, the figures of the old Irish and the improved Irish breeds in
Richardson on 'The Pig,' 1847.
[163] Quoted by Isid. Geoffroy, 'Hist. Nat. Gen.,' tom. iii. p. 441.
[164] S. Sidney, 'The Pig,' p. 61.
[165] 'Schweineschaedel,' s. 2, 20.
[166] 'Proc. Zoolog. Soc.,' 1837, p. 23. I have not given the caudal
vertebrae, as Mr. Eyton says some might possibly have been lost. I have
added together the dorsal and lumbar vertebrae, owing to Prof. Owen's
remarks ('Journal Linn. Soc.,' vol. ii. p. 28) on the difference between
dorsal and lumbar vertebrae depending only on the development of the ribs.
Nevertheless the difference in the number of the ribs in pigs deserves
notice.
[167] 'Edinburgh New Philosoph. Journal,' April 1863. _See_ also De
Blainville's 'Osteographie,' p. 128, for various authorities on this
subject.
[168] Eudes-Deslongchamps, 'Memoires de la Soc. Linn. de Normandie,' vol.
vii., 1842, p. 41. Richardson, 'Pigs, their Origin, &c.,' 1847, p. 30.
Nathusius, 'Die Racen des Schweines,' 1860, s. 54.
[169] D. Johnson's 'Sketches of Indian Field Sports,' p. 272. Mr. Crawfurd
informs me that the same fact holds good with the wild pigs of the Malay
peninsula.
[170] For Turkish pigs, _see_ Desmarest, 'Mammalogie,' 1820, p. 391. For
those of Westphalia, _see_ Richardson's 'Pigs, their Origin,' &c., 1847, p.
41.
[171] With respect to the several foregoing and following statements on
feral pigs, _see_ Roulin, in 'Mem. presentes par divers Savans a l'Acad.,'
&c., Paris, tom. vi., 1835, p. 326. It should be observed that his account
does not apply to truly feral pigs; but to pigs long introduced into the
country and living in a half-wild state. For the truly feral pigs of
Jamaica, _see_ Gosse's 'Sojourn in Jamaica,' 1851, p. 386; and Col.
Hamilton Smith, in 'Nat. Library,' vol. ix. p. 93. With respect to Africa,
_see_ Livingstone's 'Expedition to the Zambesi,' 1865, p. 153. The most
precise statement with respect to the tusks of the West Indian feral boars
is by P. Labat (quoted by Roulin); but this author attributes the state of
these pigs to descent from a domestic stock which he saw in Spain. Admiral
Sulivan, R.N., had ample opportunities of observing the wild pigs on Eagle
Islet in the Falklands; and he informs me that they resembled wild boars
with bristly ridged backs and large tusks. The pigs which have run wild in
the province of Buenos Ayres (Rengger, 'Saeugethiere,' s. 331) have not
reverted to the wild type. De Blainville ('Osteographie,' p. 132) refers to
two skulls of domestic pigs sent from Patagonia by Al. d'Orbigny, and he
states that they have the occipital elevation of the wild European boar,
but that the head altogether is "plus courte et plus ramassee." He refers,
also, to the skin of a feral pig from North America, and says, "il
ressemble tout a fait a un petit sanglier, mais il est presque tout noir,
et peut-etre un peu plus ramasse dans ses formes."
[172] Gosse's 'Jamaica,' p. 386, with a quotation from Williamson's
'Oriental Field Sports.' Also Col. Hamilton Smith, in 'Naturalist's
Library,' vol. ix. p. 94.
[173] S. Sidney's edition of 'Youatt on the Pig,' 1860, pp. 7, 26, 27, 29,
30.
[174] 'Schweineschaedel,' s. 140.
[175] 'Die Fauna der Pfahlbauten,' 1861, s. 109, 149, 222. _See_ also
Geoffroy Saint Hilaire, in 'Mem. du Mus. d'Hist. Nat.,' tom. x. p. 172; and
his son Isidore, in 'Hist. Nat. Gen.,' tom. iii. p. 69. Vasey, in his
'Delineations of the Ox Tribe,' 1851, p. 127, says the zebu has four, and
the common ox five, sacral vertebrae. Mr. Hodgson found the ribs either
thirteen or fourteen in number; _see_ a note in 'Indian Field,' 1858, p.
62.
[176] 'The Indian Field,' 1858, p. 74, where Mr. Blyth gives his
authorities with respect to the feral humped cattle. Pickering, also, in
his 'Races of Man,' 1850, p. 274, notices the peculiar character of the
grunt-like voice of the humped cattle.
[177] Mr. H. E. Marquand, in 'The Times,' June 23rd, 1856.
[178] Vasey, 'Delineations of the Ox-Tribe,' p. 124. Brace's 'Hungary,'
1851, p. 94. The Hungarian cattle descend, according to Ruetimeyer ('Zahmen.
Europ. Rindes,' 1866, s. 13), from _Bos primigenius_.
[179] Moll and Gayot, 'La Connaissance Gen. du Boeuf,' Paris, 1860. Fig 82
is that of the Podolian breed.
[180] A translation appeared in three parts in the 'Annals and Mag. of Nat.
Hist.,' 2nd series, vol. iv., 1849.
[181] _See_, also, Ruetimeyer's 'Beitrage pal. Gesch. der Wiederkauer,'
Basel, 1865, s. 54.
[182] Pictet's 'Paleontologie,' tom. i. p. 365 (2nd edit.). With respect to
B. trochoceros, _see_ Ruetimeyer's 'Zahmen Europ. Rindes,' 1866, s. 26.
[183] Owen, 'British Fossil Mammals,' 1846, p. 510.
[184] 'British Pleistocene Mammalia,' by W. B. Dawkins and W. A. Sandford,
1866. p. xv.
[185] W. R. Wilde, 'An Essay on the Animal Remains, &c., Royal Irish
Academy,' 1860, p. 29. Also 'Proc. of R. Irish Academy,' 1858, p. 48.
[186] 'Lecture: Royal Institution of G. Britain,' May 2nd, 1856, p. 4.
'British Fossil Mammals,' p. 513.
[187] Nilsson, in 'Annals and Mag. of Nat. Hist.,' 1849, vol. iv. p. 354.
[188] _See_ W. R. Wilde, ut supra; and Mr. Blyth, in 'Proc. Irish Academy,'
March 5th, 1864.
[189] Laing's 'Tour in Norway,' p. 110.
[190] Isid. Geoffroy St. Hilaire, 'Hist. Nat. Gen.,' tom. iii. p. 96.
[191] Idem, tom. iii. pp. 82, 91.
[192] 'Quadrupedes du Paraguay,' tom. ii. p. 360.
[193] Walther, 'Das Rindvieh,' 1817, s. 30.
[194] I am much indebted to the present Earl of Tankerville for information
about his wild cattle; and for the skull which was sent to Prof. Ruetimeyer.
The fullest account of the Chillingham cattle is given by Mr. Hindmarsh,
together with a letter by the late Lord Tankerville, in 'Annals and Mag. of
Nat. Hist.,' vol. ii., 1839, p. 274. _See_ Bewick, 'Quadrupeds,' 2nd edit.,
1791, p. 35, note. With respect to those of Duke of Queensberry, _see_
Pennant's 'Tour in Scotland,' p. 109. For those of Chartley, _see_ Low's
'Domesticated Animals of Britain,' 1845, p. 238. For those of Gisburne,
_see_ Bewick's 'Quadrupeds, and Encyclop. of Rural Sports,' p. 101.
[195] Boethius was born in 1470; 'Annals and Mag. of Nat. Hist.,' vol. ii.,
1839, p. 281; and vol. iv. 1849, p. 424.
[196] Youatt on Cattle, 1834, p. 48: _See_ also p. 242, on short-horn
cattle. Bell, in his 'British Quadrupeds,' p. 423, states that, after long
attending to the subject, he has found that white cattle invariably have
coloured ears.
[197] Azara, 'Des Quadrupedes du Paraguay,' tom. ii. p. 361. Azara quotes
Buffon for the feral cattle of Africa. For Texas, _see_ 'Times,' Feb. 18th,
1846.
[198] Anson's Voyage. _See_ Kerr and Porter's 'Collection,' vol. xii. p.
103.
[199] _See_ also Mr. Mackinnon's pamphlet on the Falkland Islands, p. 24.
[200] 'The Age of the Ox, Sheep, Pig,' &c., by Prof. James Simonds,
published by order of the Royal Agricult. Soc.
[201] 'Ann. Agricult. France,' April 1897. as quoted in 'The Veterinary,'
vol. xii. p. 725. I quote Tessier's observations from Youatt on Cattle, p.
527.
[202] 'The Veterinary,' vol. viii. p. 681, and vol. x. p. 268. Low's
'Domest. Animals of Great Britain,' p. 297.
[203] Mr. Ogleby, in 'Proc. Zoolog. Soc.,' 1836, p. 138, and 1840, p. 4.
[204] Leguat's Voyage, quoted by Vasey in his 'Delineations of the
Ox-tribe,' p. 132.
[205] 'Travels in South Africa,' pp. 317, 336.
[206] 'Mem. de l'Institut present. par divers Savans,' tom. vi., 1835, p.
333. For Brazil, _see_ 'Comptes Rendus,' June 15th, 1846. _See_ Azara,
'Quadrupedes du Paraguay,' tom. ii. pp. 359, 361.
[207] 'Schweineschaedel,' 1864, s. 104. Nathusius states that the form of
skull characteristic of the niata cattle occasionally appears in European
cattle; but he is mistaken, as we shall hereafter see, in supposing that
these cattle do not form a distinct race. Prof. Wyman, of Cambridge, United
States, informs me that the common cod-fish presents a similar monstrosity,
called by the fishermen the "bulldog cod." Prof. Wyman also concluded,
after making numerous inquiries in La Plata, that the niata cattle transmit
their peculiarities or form a race.
[208] Ueber Art des Zahmen Europ. Rindes, 1866, s. 28.