Chapter XXIII

DIRECT AND DEFINITE ACTION OF THE EXTERNAL CONDITIONS OF LIFE.

SLIGHT MODIFICATIONS IN PLANTS FROM THE DEFINITE ACTION OF CHANGED
CONDITIONS IN SIZE, COLOUR, CHEMICAL PROPERTIES, AND IN THE STATE OF
THE TISSUES--LOCAL DISEASES--CONSPICUOUS MODIFICATIONS FROM CHANGED
CLIMATE OR FOOD, ETC.--PLUMAGE OF BIRDS AFFECTED BY PECULIAR NUTRIMENT,
AND BY THE INOCULATION OF POISON--LAND-SHELLS--MODIFICATIONS OF ORGANIC
BEINGS IN A STATE OF NATURE THROUGH THE DEFINITE ACTION OF EXTERNAL
CONDITIONS--COMPARISON OF AMERICAN AND EUROPEAN TREES--GALLS--EFFECTS
OF PARASITIC FUNGI--CONSIDERATIONS OPPOSED TO THE BELIEF IN THE POTENT
INFLUENCE OF CHANGED EXTERNAL CONDITIONS--PARALLEL SERIES OF
VARIETIES--AMOUNT OF VARIATION DOES NOT CORRESPOND WITH THE DEGREE OF
CHANGE IN THE CONDITIONS--BUD-VARIATION--MONSTROSITIES PRODUCED BY
UNNATURAL TREATMENT--SUMMARY.

If we ask ourselves why this or that character has been modified under domestication, we are, in most cases lost in utter darkness. Many naturalists, especially of the French school, attribute every modification to the "monde ambiant," that is, to changed climate, with all its diversities of heat and cold, dampness and dryness, light and electricity, to the nature of the soil, and to varied kinds and amount of food. By the term definite action, as used in this chapter, I mean an action of such a nature that, when many individuals of the same variety are exposed during several generations to any change in their physical conditions of life, all, or nearly all the individuals, are modified in the same manner. A new sub-variety would thus be produced without the aid of selection.

I do not include under the term of definite action the effects of habit or of the increased use and disuse of various organs. Modifications of this nature, no doubt, are definitely caused by the conditions to which the beings are subjected; but they depend much less on the nature of the conditions than on the laws of growth; hence they are included under a distinct head in the {272} following chapter. We know, however, far too little of the causes and laws of variation to make a sound classification. The direct action of the conditions of life, whether leading to definite or indefinite results, is a totally distinct consideration from the effects of natural selection; for natural selection depends on the survival under various and complex circumstances of the best-fitted individuals, but has no relation whatever to the primary cause of any modification of structure.

I will first give in detail all the facts which I have been able to collect, rendering it probable that climate, food, &c., have acted so definitely and powerfully on the organisation of our domesticated productions, that they have sufficed to form new sub-varieties or races, without the aid of selection by man or of natural selection. I will then give the facts and considerations opposed to this conclusion, and finally we will weigh, as fairly as we can, the evidence on both sides.

When we reflect that distinct races of almost all our domesticated animals exist in each kingdom of Europe, and formerly even in each district of England, we are at first strongly inclined to attribute their origin to the definite action of the physical conditions of each country; and this has been the conclusion of many authors. But we should bear in mind that man annually has to choose which animals shall be preserved for breeding, and which shall be slaughtered. We have also seen that both methodical and unconscious selection were formerly practised, and are now occasionally practised by the most barbarous races, to a much greater extent than might have been anticipated. Hence it is very difficult to judge how far the difference in conditions between, for instance, the several districts in England, could have sufficed without the aid of selection to modify the breeds which have been reared in each. It may be argued that, as numerous wild animals and plants have ranged during many ages throughout Great Britain, and still retain the same character, the difference in conditions between the several districts could not have modified in so marked a manner the various native races of cattle, sheep, pigs, and horses. The same difficulty of distinguishing between selection and the definite effects of the conditions of life, is encountered in a still higher degree when we compare closely allied natural {273} forms, inhabiting two countries, such as North America and Europe, which do not differ greatly in climate, nature of soil, &c., for in this case natural selection will inevitably and rigorously have acted during a long succession of ages.

From the importance of the difficulty just alluded to, it will be
advisable to give as large a body of facts as possible, showing that
extremely slight differences in treatment, either in different parts of
the same country, or during different seasons, certainly cause an
appreciable effect, at least on varieties which are already in an
unstable condition. Ornamental flowers are good for this purpose, as
they are highly variable, and are carefully observed. All
floriculturists are unanimous that certain varieties are affected by
very slight differences in the nature of the artificial compost in
which they are grown, and by the natural soil of the district, and by
the season. Thus, a skilful judge, in writing on Carnations and
Picotees,[656] asks "where can Admiral Curzon be seen possessing the
colour, size, and strength which it has in Derbyshire? Where can
Flora's Garland be found equal to those at Slough? Where do
high-coloured flowers revel better than at Woolwich and Birmingham? Yet
in no two of these districts do the same varieties attain an equal
degree of excellence, although each may be receiving the attention of
the most skilful cultivators." The same writer then recommends every
cultivator to keep five different kinds of soil and manure, "and to
endeavour to suit the respective appetites of the plants you are
dealing with, for without such attention all hope of general success
will be vain." So it is with the Dahlia:[657] the Lady Cooper rarely
succeeds near London, but does admirably in other districts; the
reverse holds good with other varieties; and again, there are others
which succeed equally well in various situations. A skilful
gardener[658] states that he procured cuttings of an old and well-known
variety (pulchella) of Verbena, which from having been propagated in a
different situation presented a slightly different shade of colour; the
two varieties were afterwards multiplied by cuttings, being carefully
kept distinct; but in the second year they could hardly be
distinguished, and in the third year no one could distinguish them.

The nature of the season has an especial influence on certain varieties
of the Dahlia: in 1841 two varieties were pre-eminently good, and the
next year these same two were pre-eminently bad. A famous amateur[659]
asserts that in 1861 many varieties of the Rose came so untrue in
character, "that it was hardly possible to recognise them, and the
thought was not seldom entertained that the grower had lost his tally."
The same amateur[660] states that in 1862 two-thirds of his Auriculas
produced central trusses of flowers, and these are remarkable from not
keeping true; {274} and he adds that in some seasons certain varieties
of this plant all prove good, and the next season all prove bad; whilst
exactly the reverse happens with other varieties. In 1845 the editor of
the 'Gardener's Chronicle'[661] remarked how singular it was that this
year many Calceolarias tended to assume a tubular form. With
Heartsease[662] the blotched sorts do not acquire their proper
character until hot weather sets in; whilst other varieties lose their
beautiful marks as soon as this occurs.

Analogous facts have been observed with leaves: Mr. Beaton asserts[663]
that he raised at Shrubland, during six years, twenty thousand
seedlings from the Punch Pelargonium, and not one had variegated
leaves; but at Surbiton, in Surrey, one-third, or even a greater
proportion, of the seedlings from this same variety were more or less
variegated. The soil of another district in Surrey has a strong
tendency to cause variegation, as appears from information given me by
Sir F. Pollock. Verlot[664] states that the variegated strawberry
retains its character as long as grown in a dryish soil, but soon loses
it when planted in fresh and humid soil. Mr. Salter, who is well known
for his success in cultivating variegated plants, informs me that rows
of strawberries were planted in his garden in 1859, in the usual way;
and at various distances in one row, several plants simultaneously
became variegated, and what made the case more extraordinary, all were
variegated in precisely the same manner. These plants were removed, but
during the three succeeding years other plants in the same row became
variegated, and in no instance were the plants in any adjoining row
affected.

The chemical qualities, odours, and tissues of plants are often
modified by a change which seems to us slight. The Hemlock is said not
to yield conicine in Scotland. The root of the _Aconitum napellus_
becomes innocuous in frigid climates. The medicinal properties of the
Digitalis are easily affected by culture. The Rhubarb flourishes in
England, but does not produce the medicinal substance which makes the
plant so valuable in Chinese Tartary. As the _Pistacia lentiscus_ grows
abundantly in the South of France, the climate must suit it, but it
yields no mastic. The _Laurus sassafras_ in Europe loses the odour
proper to it in North America.[665] Many similar facts could be given,
and they are remarkable because it might have been thought that
definite chemical compounds would have been little liable to change
either in quality or quantity.

The wood of the American Locust-tree (_Robinia_) when grown in England
is nearly worthless, as is that of the Oak-tree when grown at the Cape
of Good Hope.[666] Hemp and flax, as I hear from Dr. Falconer, flourish
and yield plenty of seed on the plains of India, but their fibres are
brittle {275} and useless. Hemp, on the other hand, fails to produce in
England that resinous matter which is so largely used in India as an
intoxicating drug.

The fruit of the Melon is greatly influenced by slight differences in
culture and climate. Hence it is generally a better plan, according to
Naudin, to improve an old kind than to introduce a new one into any
locality. The seed of the Persian Melon produces near Paris fruit
inferior to the poorest market kinds, but at Bordeaux yields delicious
fruit.[667] Seed is annually brought from Thibet to Kashmir,[668] and
produces fruit weighing from four to ten pounds, but plants raised from
seed saved in Kashmir next year give fruit weighing only from two to
three pounds. It is well known that American varieties of the Apple
produce in their native land magnificent and brightly-coloured fruit,
but in England of poor quality and a dull colour. In Hungary there are
many varieties of the Kidney-bean, remarkable for the beauty of their
seeds, but the Rev. M. J. Berkeley[669] found that their beauty could
hardly ever be preserved in England, and in some cases the colour was
greatly changed. We have seen in the ninth chapter, with respect to
wheat, what a remarkable effect transportal from the North to the South
of France, and reversely, produced on the weight of the grain.

When man can perceive no change in plants or animals which have been exposed to a new climate or to different treatment, insects can sometimes perceive a marked change. The same species of cactus has been carried to India from Canton, Manilla, Mauritius, and from the hot-houses of Kew, and there is likewise a so-called native kind, formerly introduced from South America; all these plants are alike in appearance, but the cochineal insect flourishes only on the native kind, on which it thrives prodigiously.[670] Humboldt remarks[671] that white men "born in the torrid zone walk barefoot with impunity in the same apartment where a European, recently landed, is exposed to the attacks of the _Pulex penetrans_." This insect, the too well-known chigoe, must therefore be able to distinguish what the most delicate chemical analysis fails to distinguish, namely, a difference between the blood or tissues of a European and those of a white man born in the country. But the discernment of the chigoe is not so surprising as it at first appears; for {276} according to Liebig[672] the blood of men with different complexions, though inhabiting the same country, emits a different odour.

Diseases peculiar to certain localities, heights, or climates, may be
here briefly noticed, as showing the influence of external
circumstances on the human body. Diseases confined to certain races of
man do not concern us, for the constitution of the race may play the
more important part, and this may have been determined by unknown
causes. The Plica Polonica stands, in this respect, in a nearly
intermediate position; for it rarely affects Germans, who inhabit the
neighbourhood of the Vistula, where so many Poles are grievously
affected; and on the other hand, it does not affect Russians, who are
said to belong to the same original stock with the Poles.[673] The
elevation of a district often governs the appearance of diseases; in
Mexico the yellow fever does not extend above 924 mètres; and in Peru,
people are affected with the _verugas_ only between 600 and 1600 mètres
above the sea; many other such cases could be given. A peculiar
cutaneous complaint, called the _Bouton d'Alep_, affects in Aleppo and
some neighbouring districts almost every native infant, and some few
strangers; and it seems fairly well established that this singular
complaint depends on drinking certain waters. In the healthy little
island of St. Helena the scarlet-fever is dreaded like the Plague;
analogous facts have been observed in Chili and Mexico.[674] Even in
the different departments of France it is found that the various
infirmities which render the conscript unfit for serving in the army,
prevail with remarkable inequality, revealing, as Boudin observes, that
many of them are endemic, which otherwise would never have been
suspected.[675] Any one who will study the distribution of disease will
be struck with surprise at what slight differences in the surrounding
circumstances govern the nature and severity of the complaints by which
man is at least temporarily affected.

The modifications as yet referred to have been extremely slight, and in most cases have been caused, as far as we can judge, by equally slight changes in the conditions. But can it be safely maintained that such changed conditions, if acting during a long series of generations, would not produce a marked effect? It is commonly believed that the people of the United States differ in appearance from the parent Anglo-Saxon race; and selection cannot have come into action within so short a period. A good observer[676] states that a general absence of fat, {277} a thin and elongated neck, stiff and lank hair, are the chief characteristics. The change in the nature of the hair is supposed to be caused by the dryness of the atmosphere. If immigration into the United States were now stopped, who can say that the character of the whole people would not be greatly modified in the course of two or three thousand years?

The direct and definite action of changed conditions, in
contradistinction to the accumulation of indefinite variations, seems
to me so important that I will give a large additional body of
miscellaneous facts. With plants, a considerable change of climate
sometimes produces a conspicuous result. I have given in detail in the
ninth chapter the most remarkable case known to me, namely, that in
Germany several varieties of maize brought from the hotter parts of
America were transformed in the course of only two or three
generations. Dr. Falconer informs me that he has seen the English
Ribston-pippin apple, a Himalayan oak, Prunus and Pyrus, all assume in
the hotter parts of India a fastigate or pyramidal habit; and this fact
is the more interesting, as a Chinese tropical species of Pyrus
naturally has this habit of growth. Although in these cases the changed
manner of growth seems to have been directly caused by the great heat,
we know that many fastigate trees have originated in their temperate
homes. In the Botanic Gardens of Ceylon the apple-tree[677] "sends out
numerous runners under ground, which continually rise into small stems,
and form a growth around the parent-tree." The varieties of the cabbage
which produce heads in Europe fail to do so in certain tropical
countries.[678] The _Rhododendron ciliatum_ produced at Kew flowers so
much larger and paler-coloured than those which it bears on its native
Himalayan mountain, that Dr. Hooker[679] would hardly have recognised
the species by the flowers alone. Many similar facts with respect to
the colour and size of flowers could be given.

The experiments of Vilmorin and Buckman on carrots and parsnips prove
that abundant nutriment produces a definite and inheritable effect on
the so-called roots, with scarcely any change in other parts of the
plant. Alum directly influences the colour of the flowers of the
Hydrangea.[680] Dryness seems generally to favour the hairyness or
villosity of plants. Gärtner found that hybrid Verbascums became
extremely woolly when grown in pots. Mr. Masters, on the other hand,
states that the _Opuntia leucotricha_ "is well clothed with beautiful
white hairs when grown in a damp heat; but in a dry heat exhibits none
of this peculiarity."[681] Slight variations of many kinds, not worth
specifying in detail, are retained only as {278} long as plants are
grown in certain soils, of which Sageret[682] gives from his own
experience some instances. Odart, who insists strongly on the
permanence of the varieties of the grape, admits[683] that some
varieties, when grown under a different climate or treated differently,
vary in an extremely slight degree, as in the tint of the fruit and in
the period of ripening. Some authors have denied that grafting causes
even the slightest difference in the scion; but there is sufficient
evidence that the fruit is sometimes slightly affected in size and
flavour, the leaves in duration, and the flowers in appearance.[684]

With animals there can be no doubt, from the facts given in the first
chapter, that European dogs deteriorate in India, not only in their
instincts but in structure; but the changes which they undergo are of
such a nature, that they may be partly due to reversion to a primitive
form, as in the case of feral animals. In parts of India the turkey
becomes reduced in size, "with the pendulous appendage over the beak
enormously developed."[685] We have seen how soon the wild duck, when
domesticated, loses its true character, from the effects of abundant or
changed food, or from taking little exercise. From the direct action of
a humid climate and poor pasture the horse rapidly decreases in size in
the Falkland Islands. From information which I have received, this
seems likewise to be the case to a certain extent with sheep in
Australia.

Climate definitely influences the hairy covering of animals; in the
West Indies a great change is produced in the fleece of sheep, in about
three generations. Dr. Falconer states[686] that the Thibet mastiff and
goat, when brought down from the Himalaya to Kashmir, lose their fine
wool. At Angora not only goats, but shepherd-dogs and cats, have fine
fleecy hair, and Mr. Ainsworth[687] attributes the thickness of the
fleece to the severe winters, and its silky lustre to the hot summers.
Burnes states positively[688] that the Karakool sheep lose their
peculiar black curled fleeces when removed into any other country. Even
within the limits of England, I have been assured that with two breeds
of sheep the wool was slightly changed by the flocks being pastured in
different localities.[689] It has been asserted on good authority[690]
that horses kept during several years in the deep coal-mines of Belgium
become covered with velvety hair, almost like that on the mole. These
cases probably stand in close relation to the natural change of coat in
winter and summer. Naked varieties of several domestic animals have
occasionally appeared; but there is no reason to {279} believe that
this is in any way related to the nature of the climate to which they
have been exposed.[691]

It appears at first sight probable that the increased size, the
tendency to fatten, the early maturity and altered forms of our
improved cattle, sheep, and pigs, have directly resulted from their
abundant supply of food. This is the opinion of many competent judges,
and probably is to a great extent true. But as far as form is
concerned, we must not overlook the equal or more potent influence of
lessened use on the limbs and lungs. We see, moreover, as far as size
is concerned, that selection is apparently a more powerful agent than a
large supply of food, for we can thus only account for the existence,
as remarked to me by Mr. Blyth, of the largest and smallest breeds of
sheep in the same country, of Cochin-China fowls and Bantams, of small
Tumbler and large Runt pigeons, all kept together and supplied with
abundant nourishment. Nevertheless there can be little doubt that our
domesticated animals have been modified, independently of the increased
or lessened use of parts, by the conditions to which they have been
subjected, without the aid of selection. For instance, Prof.
Rütimeyer[692] shows that the bones of all domesticated quadrupeds can
be distinguished from those of wild animals by the state of their
surface and general appearance. It is scarcely possible to read
Nathusius's excellent 'Vorstudien,'[693] and doubt that, with the
highly improved races of the pig, abundant food has produced a
conspicuous effect on the general form of the body, on the breadth of
the head and face, and even on the teeth. Nathusius rests much on the
case of a purely bred Berkshire pig, which when two months old became
diseased in its digestive organs, and was preserved for observation
until nineteen months old; at this age it had lost several
characteristic features of the breed, and had acquired a long, narrow
head, of large size relatively to its small body, and elongated legs.
But in this case and in some others we ought not to assume that,
because certain characters are lost, perhaps through reversion, under
one course of treatment, therefore that they had been at first directly
produced by an opposite course.

In the case of the rabbit, which has become feral on the island of
Porto Santo, we are at first strongly tempted to attribute the whole
change--the greatly reduced size, the altered tints of the fur, and the
loss of certain characteristic marks--to the definite action of the new
conditions to which it has been exposed. But in all such cases we have
to consider in addition the tendency to reversion to progenitors more
or less remote, and the natural selection of the finest shades of
difference.

The nature of the food sometimes either definitely induces certain
peculiarities, or stands in some close relation with them. Pallas long
ago asserted that the fat-tailed sheep of Siberia degenerated and lost
their enormous tails when removed from certain saline pastures; and
recently {280} Erman[694] states that this occurs with the Kirgisian
sheep when brought to Orenburgh.

It is well known that hemp-seed causes bullfinches and certain other
birds to become black. Mr. Wallace has communicated to me some much
more remarkable facts of the same nature. The natives of the Amazonian
region feed the common green parrot (_Chrysotis festiva_, Linn.) with
the fat of large Siluroid fishes, and the birds thus treated become
beautifully variegated with red and yellow feathers. In the Malayan
archipelago, the natives of Gilolo alter in an analogous manner the
colours of another parrot, namely, the _Lorius garrulus_, Linn., and
thus produce the _Lori rajah_ or King-Lory. These parrots in the Malay
Islands and South America, when fed by the natives on natural vegetable
food, such as rice and plantains, retain their proper colours. Mr.
Wallace has, also, recorded[695] a still more singular fact. "The
Indians (of S. America) have a curious art by which they change the
colours of the feathers of many birds. They pluck out those from the
part they wish to paint, and inoculate the fresh wound with the milky
secretion from the skin of a small toad. The feathers grow of a
brilliant yellow colour, and on being plucked out, it is said, grow
again of the same colour without any fresh operation."

Bechstein[696] does not entertain any doubt that seclusion from light
affects, at least temporarily, the colours of cage-birds.

It is well known that the shells of land-mollusca are affected by the
abundance of lime in different districts. Isidore Geoffroy St.
Hilaire[697] gives the case of _Helix lactea_, which has recently been
carried from Spain to the South of France and to the Rio Plata, and in
both these countries now presents a distinct appearance, but whether
this has resulted from food or climate is not known. With respect to
the common oyster, Mr. F. Buckland informs me that he can generally
distinguish the shells from different districts; young oysters brought
from Wales and laid down in beds where "_natives_" are indigenous, in
the short space of two months begin to assume the "native" character.
M. Costa[698] has recorded a much more remarkable case of the same
nature, namely, that young shells taken from the shores of England and
placed in the Mediterranean, at once altered their manner of growth and
formed prominent diverging rays, like those on the shells of the proper
Mediterranean oyster. The same individual shell, showing both forms of
growth, was exhibited before a society in Paris. Lastly, it is well
known that caterpillars fed on different food sometimes either
themselves acquire a different colour or produce moths different in
colour.[699]

{281}

It would be travelling beyond my proper limits here to discuss how far
organic beings in a state of nature are definitely modified by changed
conditions. In my 'Origin of Species' I have given a brief abstract of
the facts bearing on this point, and have shown the influence of light
on the colours of birds, and of residence near the sea on the lurid
tints of insects, and on the succulency of plants. Mr. Herbert
Spencer[700] has recently discussed with much ability this whole
subject on broad and general grounds. He argues, for instance, that
with all animals the external and internal tissues are differently
acted on by the surrounding conditions, and they invariably differ in
intimate structure. So again the upper and lower surfaces of true
leaves, as well as of stems and petioles, when these assume the
function and occupy the position of leaves, are differently
circumstanced with respect to light, &c., and apparently in consequence
differ in structure. But, as Mr. Herbert Spencer admits, it is most
difficult in all such cases to distinguish between the effects of the
definite action of physical conditions and the accumulation through
natural selection of inherited variations which are serviceable to the
organism, and which have arisen independently of the definite action of
these conditions.

Although we are not here concerned with organic beings in a state of nature, yet I may call attention to one case. Mr. Meehan,[701] in a remarkable paper, compares twenty-nine kinds of American trees, belonging to various orders, with their nearest European allies, all grown in close proximity in the same garden and under as nearly as possible the same conditions. In the American species Mr. Meehan finds, with the rarest exceptions, that the leaves fall earlier in the season, and assume before falling a brighter tint; that they are less deeply toothed or serrated; that the buds are smaller; that the trees are more diffuse in growth and have fewer branchlets; and, lastly, that the seeds are smaller--all in comparison with the corresponding European species. Now, considering that these trees belong to distinct orders, it is out of the question that the peculiarities just specified should have been inherited in the one continent from one progenitor, and in the other from another progenitor; and considering that the trees inhabit widely different stations, these peculiarities can hardly be supposed to be of any special {282} service to the two series in the Old and New Worlds; therefore these peculiarities cannot have been naturally selected. Hence we are led to infer that they have been definitely caused by the long-continued action of the different climate of the two continents on the trees.

_Galls._--Another class of facts, not relating to cultivated plants, deserves attention. I allude to the production of galls. Every one knows the curious, bright-red, hairy productions on the wild rose-tree, and the various different galls produced by the oak. Some of the latter resemble fruit, with one face as rosy as the rosiest apple. These bright colours can be of no service either to the gall-forming insect or to the tree, and probably are the direct result of the action of the light, in the same manner as the apples of Nova Scotia or Canada are brighter coloured than English apples. The strongest upholder of the doctrine that organic beings are created beautiful to please mankind would not, I presume, extend this view to galls. According to Osten Sacken's latest revision, no less than fifty-eight kinds of galls are produced on the several species of oak, by Cynips with its sub-genera; and Mr. B. D. Walsh[702] states that he can add many others to the list. One American species of willow, the _Salix humilis_, bears ten distinct kinds of galls. The leaves which spring from the galls of various English willows differ completely in shape from the natural leaves. The young shoots of junipers and firs, when punctured by certain insects, yield monstrous growths like flowers and cones; and the flowers of some plants become from the same cause wholly changed in appearance. Galls are produced in every quarter of the world; of several sent to me by Mr. Thwaites from Ceylon, some were as symmetrical as a composite flower when in bud, others smooth and spherical like a berry; some protected by long spines, others clothed with yellow wool formed of long cellular hairs, others with regularly tufted hairs. In some galls the internal structure is simple, but in others it is highly complex; thus M. Lucaze-Duthiers[703] has figured in the common ink-gall no less than seven concentric layers, composed of distinct tissue, {283} namely, the epidermic, sub-epidermic, spongy, intermediate, and the hard protective layer formed of curiously thickened woody cells, and, lastly, the central mass abounding with starch-granules on which the larvæ feed.

Galls are produced by insects of various orders, but the greater number by species of Cynips. It is impossible to read M. Lucaze-Duthier's discussion and doubt that the poisonous secretion of the insect causes the growth of the gall, and every one knows how virulent is the poison secreted by wasps and bees, which belong to the same order with Cynips. Galls grow with extraordinary rapidity, and it is said that they attain their full size in a few days;[704] it is certain that they are almost completely developed before the larvæ are hatched. Considering that many gall-insects are extremely small, the drop of secreted poison must be excessively minute; it probably acts on one or two cells alone, which, being abnormally stimulated, rapidly increase by a process of self-division. Galls, as Mr. Walsh[705] remarks, afford good, constant, and definite characters, each kind keeping as true to form as does any independent organic being. This fact becomes still more remarkable when we hear that, for instance, seven out of the ten different kinds of galls produced on _Salix humilis_ are formed by gall-gnats (_Cecidomyidæ_) which, "though essentially distinct species, yet resemble one another so closely that in almost all cases it is difficult, and in some cases impossible, to distinguish the full-grown insects one from the other."[706] For in accordance with a wide-spread analogy we may safely infer that the poison secreted by insects so closely allied would not differ much in nature; yet this slight difference is sufficient to induce widely different results. In some few cases the same species of gall-gnat produces on distinct species of willows galls which cannot be distinguished; the _Cynips fecundatrix_, also, has been known to produce on the Turkish oak, to which it is not properly attached, exactly the same kind of gall as on the European oak.[707] These latter facts apparently prove that the nature of the poison is a much more powerful {284} agent in determining the form of the gall than the specific character of the tree which is acted on.

As the poisonous secretion of insects belonging to various orders has the special power of affecting the growth of various plants;--as a slight difference in the nature of the poison suffices to produce widely different results;--and lastly, as we know that the chemical compounds secreted by plants are eminently liable to be modified by changed conditions of life, we may believe it possible that various parts of a plant might be modified through the agency of its own altered secretions. Compare, for instance, the mossy and viscid calyx of a moss-rose, which suddenly appears through bud-variation on a Provence-rose, with the gall of red moss growing from the inoculated leaf of a wild rose, with each filament symmetrically branched like a microscopical spruce-fir, bearing a glandular tip and secreting odoriferous gummy matter.[708] Or compare, on the one hand, the fruit of the peach, with its hairy skin, fleshy covering, hard shell and kernel, and on the other hand one of the more complex galls with its epidermic, spongy, and woody layers, surrounding tissue loaded with starch granules. These normal and abnormal structures manifestly present a certain degree of resemblance. Or, again, reflect on the cases above given of parrots which have had their plumage brightly decorated through some change in their blood, caused by having been fed on certain fishes, or locally inoculated with the poison of a toad. I am far from wishing to maintain that the moss-rose or the hard shell of the peach-stone or the bright colours of birds are actually due to any chemical change in the sap or blood; but these cases of galls and of parrots are excellently adapted to show us how powerfully and singularly external agencies may affect structure. With such facts before us, we need feel no surprise at the appearance of any modification in any organic being.

I may, also, here allude to the remarkable effects which parasitic
fungi sometimes produce on plants. Reissek[709] has described a
Thesium, affected by an Oecidium, which was greatly modified, and
assumed some of the {285} characteristic features of certain allied
species, or even genera. Suppose, says Reissek, "the condition
originally caused by the fungus to become constant in the course of
time, the plant would, if found growing wild, be considered as a
distinct species or even as belonging to a new genus." I quote this
remark to show how profoundly, yet in how natural a manner, this plant
must have been modified by the parasitic fungus.

_Facts and Considerations opposed to the belief that the Conditions of Life act in a potent manner in causing definite Modifications of Structure._

I have alluded to the slight differences in species when naturally living in distinct countries under different conditions; and such differences we feel at first inclined, probably to a limited extent with justice, to attribute to the definite action of the surrounding conditions. But it must be borne in mind that there are a far greater number of animals and plants which range widely and have been exposed to great diversities of conditions, yet remain nearly uniform in character. Some authors, as previously remarked, account for the varieties of our culinary and agricultural plants by the definite action of the conditions to which they have been exposed in the different parts of Great Britain; but there are about 200 plants[710] which are found in every single English county; these plants must have been exposed for an immense period to considerable differences of climate and soil, yet do not differ. So, again, some birds, insects, other animals, and plants range over large portions of the world, yet retain the same character.

Notwithstanding the facts previously given on the occurrence of highly
peculiar local diseases and on the strange modifications of structure
in plants caused by the inoculated poison of insects, and other
analogous cases; still there are a multitude of variations--such as the
modified skull of the niata ox and bulldog, the long horns of Caffre
cattle, the conjoined toes of the solid-hoofed swine, the immense crest
and protuberant skull of Polish fowls, the crop of the pouter-pigeon,
and a host of other such cases--which we can hardly attribute to the
definite action, in the sense before specified, of the external
conditions of life. No doubt in every case there must have been some
exciting cause; but as we see innumerable individuals exposed to nearly
the same conditions, and one alone is affected, we may conclude that
the constitution of the individual is of far higher {286} importance
than the conditions to which it has been exposed. It seems, indeed, to
be a general rule that conspicuous variations occur rarely, and in one
individual alone out of many thousands, though all may have been
exposed, as far as we can judge, to nearly the same conditions. As the
most strongly marked variations graduate insensibly into the most
trifling, we are led by the same train of thought to attribute each
slight variation much more to innate differences of constitution,
however caused, than to the definite action of the surrounding
conditions.

We are led to the same conclusion by considering the cases, formerly
alluded to, of fowls and pigeons, which have varied and will no doubt
go on varying in directly opposite ways, though kept during many
generations under nearly the same conditions. Some, for instance, are
born with their beaks, wings, tails, legs, &c., a little longer, and
others with these same parts a little shorter. By the long-continued
selection of such slight individual differences, which occur in birds
kept in the same aviary, widely different races could certainly be
formed; and long-continued selection, important as is the result, does
nothing but preserve the variations which appear to us to arise
spontaneously.

In these cases we see that domesticated animals vary in an indefinite
number of particulars, though treated as uniformly as is possible. On
the other hand, there are instances of animals and plants, which,
though exposed to very different conditions, both under nature and
domestication, have varied in nearly the same manner. Mr. Layard
informs me that he has observed amongst the Caffres of South Africa a
dog singularly like an arctic Esquimaux dog. Pigeons in India present
nearly the same wide diversities of colour as in Europe; and I have
seen chequered and simply barred pigeons, and pigeons with blue and
white loins, from Sierra Leone, Madeira, England, and India. New
varieties of flowers are continually raised in different parts of Great
Britain, but many of these are found by the judges at our exhibitions
to be almost identical with old varieties. A vast number of new
fruit-trees and culinary vegetables have been produced in North
America: these differ from European varieties in the same general
manner as the several varieties raised in Europe differ from each
other; and no one has ever pretended that the climate of America has
given to the many American varieties any general character by which
they can be recognised. Nevertheless, from the facts previously
advanced on the authority of Mr. Meehan with respect to American and
European forest-trees, it would be rash to affirm that varieties raised
in the two countries would not in the course of ages assume a
distinctive character. Mr. Masters has recorded a striking fact[711]
bearing on this subject: he raised numerous plants of _Hybiscus
Syriacus_ from seed collected in South Carolina and the Holy Land,
where the parent-plants must have been exposed to considerably
different conditions; yet the seedlings from both localities broke into
two similar strains, one with obtuse leaves and purple or crimson
flowers, and the other with elongated leaves and more or less pink
flowers.

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We may, also, infer the prepotent influence of the constitution of the
organism over the definite action of the conditions of life, from the
several cases given in the earlier chapters of parallel series of
varieties,--an important subject, hereafter to be more fully discussed.
Sub-varieties of the several kinds of wheat, gourds, peaches, and other
plants, and to a certain limited extent sub-varieties of the fowl,
pigeon, and dog, have been shown either to resemble or to differ from
each other in a closely corresponding and parallel manner. In other
cases, a variety of one species resembles a distinct species; or the
varieties of two distinct species resemble each other. Although these
parallel resemblances no doubt often result from reversion to the
former characters of a common progenitor; yet in other cases, when new
characters first appear, the resemblance must be attributed to the
inheritance of a similar constitution, and consequently to a tendency
to vary in the same manner. We see something of a similar kind in the
same monstrosity appearing and reappearing many times in the same
animal, and, as Dr. Maxwell Masters has remarked to me, in the same
plant.

We may at least conclude thus far, that the amount of modification which animals and plants have undergone under domestication, does not correspond with the degree to which they have been subjected to changed circumstances. As we know the parentage of domesticated birds far better than of most quadrupeds, we will glance through the list. The pigeon has varied in Europe more than almost any other bird; yet it is a native species, and has not been exposed to any extraordinary change of conditions. The fowl has varied equally, or almost equally, with the pigeon, and is a native of the hot jungles of India. Neither the peacock, a native of the same country, nor the guinea-fowl, an inhabitant of the dry deserts of Africa, has varied at all, or only in colour. The turkey, from Mexico, has varied but little. The duck, on the other hand, a native of Europe, has yielded some well-marked races; and as this is an aquatic bird, it must have been subjected to a far more serious change in its habits than the pigeon or even the fowl, which nevertheless have varied in a much higher degree. The goose, a native of Europe and aquatic like the duck, has varied less than any other domesticated bird, except the peacock.

Bud-variation is, also, important under our present point of view. In some few cases, as when all the eyes or buds on the same tuber of the potato, or all the fruit on the same plum-tree, or all the flowers on the same plant, have suddenly varied in the same manner, it might be argued that the {288} variation had been definitely caused by some change in the conditions to which the plants had been exposed; yet, in other cases, such an admission is extremely difficult. As new characters sometimes appear by bud-variation, which do not occur in the parent-species or in any allied species, we may reject, at least in these cases, the idea that they are due to reversion. Now it is well worth while to reflect maturely on some striking case of bud-variation, for instance that of the peach. This tree has been cultivated by the million in various parts of the world, has been treated differently, grown on its own roots and grafted on various stocks, planted as a standard, against a wall, and under glass; yet each bud of each sub-variety keeps true to its kind. But occasionally, at long intervals of time, a tree in England, or under the widely-different climate of Virginia, produces a single bud, and this yields a branch which ever afterwards bears nectarines. Nectarines differ, as every one knows, from peaches in their smoothness, size, and flavour; and the difference is so great, that some botanists have maintained that they are specifically distinct. So permanent are the characters thus suddenly acquired, that a nectarine produced by bud-variation has propagated itself by seed. To guard against the supposition that there is some fundamental distinction between bud and seminal variation, it is well to bear in mind that nectarines have likewise been produced from the stone of the peach; and, reversely, peaches from the stone of the nectarine. Now is it possible to conceive external conditions more closely alike than those to which the buds on the same tree are exposed? Yet one bud alone, out of the many thousands borne by the same tree, has suddenly without any apparent cause produced a nectarine. But the case is even stronger than this, for the same flower-bud has yielded a fruit, one-half or one-quarter a nectarine, and the other half or three-quarters a peach. Again, seven or eight varieties of the peach have yielded by bud-variation nectarines: the nectarines thus produced, no doubt, differ a little from each other; but still they are nectarines. Of course there must be some cause, internal or external, to excite the peach-bud to change its nature; but I cannot imagine a class of facts better adapted to force on our minds the conviction that what we call the external conditions of life are quite insignificant in {289} relation to any particular variation, in comparison with the organisation or constitution of the being which varies.

It is known from the labours of Geoffroy St. Hilaire, and recently from those of Dareste and others, that eggs of the fowl, if shaken, placed upright, perforated, covered in part with varnish, &c., produce monstrous chickens. Now these monstrosities may be said to be directly caused by such unnatural conditions, but the modifications thus induced are not of a definite nature. An excellent observer, M. Camille Dareste,[712] remarks "that the various species of monstrosities are not determined by specific causes; the external agencies which modify the development of the embryo act solely in causing a perturbation--a perversion in the normal course of development." He compares the result to what we see in illness: a sudden chill, for instance, affects one individual alone out of many, causing either a cold, or sore-throat, rheumatism, or inflammation of the lungs or pleura. Contagious matter acts in an analogous manner.[713] We may take a still more specific instance: seven pigeons were struck by rattle-snakes;[714] some suffered from convulsions; some had their blood coagulated, in others it was perfectly fluid; some showed ecchymosed spots on the heart, others on the intestines, &c.; others again showed no visible lesion in any organ. It is well known that excess in drinking causes different diseases in different men; but men living under a cold and tropical climate are differently affected:[715] and in this case we see the definite influence of opposite conditions. The foregoing facts apparently give us as good an idea as we are likely for a long time to obtain, how in many cases external conditions act directly, though not definitely, in causing modifications of structure.

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_Summary._--There can be no doubt, from the facts given in the early part of this chapter, that extremely slight changes in {290} the conditions of life sometimes act in a definite manner on our already variable domesticated productions; and, as the action of changed conditions in causing general or indefinite variability is accumulative, so it may be with their definite action. Hence it is possible that great and definite modifications of structure may result from altered conditions acting during a long series of generations. In some few instances a marked effect has been produced quickly on all, or nearly all, the individuals which have been exposed to some considerable change of climate, food, or other circumstance. This has occurred, and is now occurring, with European men in the United States, with European dogs in India, with horses in the Falkland Islands, apparently with various animals at Angora, with foreign oysters in the Mediterranean, and with maize grown in Europe from tropical seed. We have seen that the chemical compounds secreted by plants and the state of their tissues are readily affected by changed conditions. In some cases a relation apparently exists between certain characters and certain conditions, so that if the latter be changed the character is lost--as with cultivated flowers, with some few culinary plants, with the fruit of the melon, with fat-tailed sheep, and other sheep having peculiar fleeces.

The production of galls, and the change of plumage in parrots when fed on peculiar food or when inoculated by the poison of a toad, prove to us what great and mysterious changes in structure and colour may be the definite result of chemical changes in the nutrient fluids or tissues.

We have also reason to believe that organic beings in a state of nature may be modified in various definite ways by the conditions to which they have been long exposed, as in the case of American trees in comparison with their representatives in Europe. But in all such cases it is most difficult to distinguish between the definite results of changed conditions, and the accumulation through natural selection of serviceable variations which have arisen independently of the nature of the conditions. If, for instance, a plant had to be modified so as to become fitted to inhabit a humid instead of an arid station, we have no reason to believe that variations of the right kind would occur more frequently if the parent-plant inhabited a station a little more {291} humid than usual. Whether the station was unusually dry or humid, variations adapting the plant in a slight degree for directly opposite habits of life would occasionally arise, as we have reason to believe from what we know in other cases.

In most, perhaps in all cases, the organisation or constitution of the being which is acted on, is a much more important element than the nature of the changed conditions, in determining the nature of the variation. We have evidence of this in the appearance of nearly similar modifications under different conditions, and of different modifications under apparently nearly the same conditions. We have still better evidence of this in closely parallel varieties being frequently produced from distinct races, or even distinct species, and in the frequent recurrence of the same monstrosity in the same species. We have also seen that the degree to which domesticated birds have varied, does not stand in any close relation with the amount of change to which they have been subjected.

To recur once again to bud-variations. When we reflect on the millions of buds which many trees have produced, before some one bud has varied, we are lost in wonder what the precise cause of each variation can be. Let us recall the case given by Andrew Knight of the forty-year-old tree of the yellow magnum bonum plum, an old variety which has been propagated by grafts on various stocks for a very long period throughout Europe and North America, and on which a single bud suddenly produced the red magnum bonum. We should also bear in mind that distinct varieties, and even distinct species,--as in the case of peaches, nectarines, and apricots,--of certain roses and camellias,--although separated by a vast number of generations from any progenitor in common, and although cultivated under diversified conditions, have yielded by bud-variation closely analogous varieties. When we reflect on these facts we become deeply impressed with the conviction that in such cases the nature of the variation depends but little on the conditions to which the plant has been exposed, and not in any especial manner on its individual character, but much more on the general nature or constitution, inherited from some remote progenitor, of the whole group of allied beings to which the plant belongs. We are thus driven to conclude that in most {292} cases the conditions of life play a subordinate part in causing any particular modification; like that which a spark plays, when a mass of combustibles bursts into flame--the nature of the flame depending on the combustible matter, and not on the spark.

No doubt each slight variation must have its efficient cause; but it is as hopeless an attempt to discover the cause of each as to say why a chill or a poison affects one man differently from another. Even with modifications resulting from the definite action of the conditions of life, when all or nearly all the individuals, which have been similarly exposed, are similarly affected, we can rarely see the precise relation between cause and effect. In the next chapter it will be shown that the increased use or disuse of various organs, produces an inherited effect. It will further be seen that certain variations are bound together by correlation and other laws. Beyond this we cannot at present explain either the causes or manner of action of Variation.

Finally, as indefinite and almost illimitable variability is the usual result of domestication and cultivation, with the same part or organ varying in different individuals in different or even in directly opposite ways; and as the same variation, if strongly pronounced, usually recurs only after long intervals of time, any particular variation would generally be lost by crossing, reversion, and the accidental destruction of the varying individuals, unless carefully preserved by man. Hence, although it must be admitted that new conditions of life do sometimes definitely affect organic beings, it may be doubted whether well-marked races have often been produced by the direct action of changed conditions without the aid of selection either by man or nature.

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