INHERITANCE continued.--FIXEDNESS OF CHARACTER--PREPOTENCY--SEXUAL LIMITATION
--CORRESPONDENCE OF AGE.
FIXEDNESS OF CHARACTER APPARENTLY NOT DUE TO ANTIQUITY OF INHERITANCE.
PREPOTENCY OF TRANSMISSION IN INDIVIDUALS OF THE SAME FAMILY, IN CROSSED
BREEDS AND SPECIES; OFTEN STRONGER IN ONE SEX THAN THE OTHER; SOMETIMES DUE TO
THE SAME CHARACTER BEING PRESENT AND VISIBLE IN ONE BREED AND LATENT IN THE
OTHER.
INHERITANCE AS LIMITED BY SEX.
NEWLY-ACQUIRED CHARACTERS IN OUR DOMESTICATED ANIMALS OFTEN TRANSMITTED BY ONE
SEX ALONE, SOMETIMES LOST BY ONE SEX ALONE.
INHERITANCE AT CORRESPONDING PERIODS OF LIFE.
THE IMPORTANCE OF THE PRINCIPLE WITH RESPECT TO EMBRYOLOGY; AS EXHIBITED IN
DOMESTICATED ANIMALS: AS EXHIBITED IN THE APPEARANCE AND DISAPPEARANCE OF
INHERITED DISEASES; SOMETIMES SUPERVENING EARLIER IN THE CHILD THAN IN
THE PARENT.
SUMMARY OF THE THREE PRECEDING CHAPTERS.
In the last two chapters the nature and force of Inheritance, the
circumstances which interfere with its power, and the tendency to Reversion,
with its many remarkable contingencies, were discussed. In the present chapter
some other related phenomena will be treated of, as fully as my materials
permit.
FIXEDNESS OF CHARACTER.
It is a general belief amongst breeders that the longer any character has been
transmitted by a breed, the more fully it will continue to be transmitted. I
do not wish to dispute the truth of the proposition that inheritance gains
strength simply through long continuance, but I doubt whether it can be
proved. In one sense the proposition is little better than a truism; if any
character has remained constant during many generations, it will be likely to
continue so, if the conditions of life remain the same. So, again, in
improving a breed, if care be taken for a length of time to exclude all
inferior individuals, the breed will obviously tend to become truer, as it
will not have been crossed during many generations by an inferior animal. We
have previously seen, but without being able to assign any cause, that, when a
new character appears, it is occasionally from the first constant, or
fluctuates much, or wholly fails to be transmitted. So it is with the
aggregate of slight differences which characterise a new variety, for some
propagate their kind from the first much truer than others. Even with plants
multiplied by bulbs, layers, etc., which may in one sense be said to form
parts of the same individual, it is well known that certain varieties retain
and transmit through successive bud-generations their newly-acquired
characters more truly than others. In none of these, nor in the following
cases, does there appear to be any relation between the force with which a
character is transmitted and the length of time during which it has been
transmitted. Some varieties, such as white and yellow hyacinths and white
sweet-peas, transmit their colours more faithfully than do the varieties which
have retained their natural colour. In the Irish family, mentioned in the
twelfth chapter, the peculiar tortoiseshell-like colouring of the eyes was
transmitted far more faithfully than any ordinary colour. Ancon and Mauchamp
sheep and niata cattle, which are all comparatively modern breeds, exhibit
remarkably strong powers of inheritance. Many similar cases could be adduced.
As all domesticated animals and cultivated plants have varied, and yet are
descended from aboriginally wild forms, which no doubt had retained the same
character from an immensely remote epoch, we see that scarcely any degree of
antiquity ensures a character being transmitted perfectly true. In this case,
however, it may be said that changed conditions of life induce certain
modifications, and not that the power of inheritance fails; but in every case
of failure, some cause, either internal or external, must interfere. It will
generally be found that the organs or parts which in our domesticated
productions have varied, or which still continue to vary,--that is, which fail
to retain their former state,--are the same with the parts which differ in the
natural species of the same genus. As, on the theory of descent with
modification, the species of the same genus have been modified since they
branched off from a common progenitor, it follows that the characters by which
they differ from one another have varied, whilst other parts of the
organisation have remained unchanged; and it might be argued that these same
characters now vary under domestication, or fail to be inherited, from their
lesser antiquity. But variation in a state of nature seems to stand in some
close relation with changed conditions of life, and characters which have
already varied under such conditions would be apt to vary under the still
greater changes consequent on domestication, independently of their greater or
less antiquity.
Fixedness of character, or the strength of inheritance, has often been judged
of by the preponderance of certain characters in the crossed offspring between
distinct races; but prepotency of transmission here comes into play, and this,
as we shall immediately see, is a very different consideration from the
strength or weakness of inheritance. (14/1. See 'Youatt on Cattle' pages 92,
69, 78, 88, 163; and 'Youatt on Sheep' page 325. Also Dr. Lucas 'L'Hered.
Nat.' tome 2 page 310.) It has often been observed that breeds of animals
inhabiting wild and mountainous countries cannot be permanently modified by
our improved breeds; and as these latter are of modern origin, it has been
thought that the greater antiquity of the wilder breeds has been the cause of
their resistance to improvement by crossing; but it is more probably due to
their structure and constitution being better adapted to the surrounding
conditions. When plants are first subjected to culture, it has been found
that, during several generations, they transmit their characters truly, that
is, do not vary, and this has been attributed to ancient characters being
strongly inherited: but it may with equal or greater probability be consequent
on changed conditions of life requiring a long time for their cumulative
action. Notwithstanding these considerations, it would perhaps be rash to deny
that characters become more strongly fixed the longer they are transmitted;
but I believe that the proposition resolves itself into this,--that characters
of all kinds, whether new or old, tend to be inherited, and that those which
have already withstood all counteracting influences and been truly
transmitted, will, as a general rule, continue to withstand them, and
consequently be faithfully inherited.
PREPOTENCY IN THE TRANSMISSION OF CHARACTER.
When individuals, belonging to the same family, but distinct enough to be
recognised, or when two well-marked races, or two species, are crossed, the
usual result, as stated in the previous chapter, is, that the offspring in the
first generation are intermediate between their parents, or resemble one
parent in one part and the other parent in another part. But this is by no
means the invariable rule; for in many cases it is found that certain
individuals, races, and species, are prepotent in transmitting their likeness.
This subject has been ably discussed by Prosper Lucas (14/2. 'Hered. Nat.'
tome 2 pages 112-120.), but is rendered extremely complex by the prepotency
sometimes running equally in both sexes, and sometimes more strongly in one
sex than in the other; it is likewise complicated by the presence of secondary
sexual characters, which render the comparison of crossed breeds with their
parents difficult.
It would appear that in certain families some one ancestor, and after him
others in the same family, have had great power in transmitting their likeness
through the male line; for we cannot otherwise understand how the same
features should so often be transmitted after marriages with many females, as
in the case of the Austrian Emperors; and so it was, according to Niebuhr,
with the mental qualities of certain Roman families. (14/3. Sir H. Holland
'Chapters on Mental Physiology' 1852 page 234.) The famous bull Favourite is
believed (14/4. 'Gardener's Chronicle' 1860 page 270.) to have had a prepotent
influence on the shorthorn race. It has also been observed (14/5. Mr. N.H.
Smith 'Observations on Breeding' quoted in 'Encyclop. of Rural Sports' page
278.) with English racehorses that certain mares have generally transmitted
their own character, whilst other mares of equally pure blood have allowed the
character of the sire to prevail. A famous black greyhound, Bedlamite, as I
hear from Mr. C.M. Brown "invariably got all his puppies black, no matter what
was the colour of the bitch;" but then Bedlamite "had a preponderance of black
in his blood, both on the sire and dam side."
[The truth of the principle of prepotency comes out more clearly when distinct
races are crossed. The improved Shorthorns, notwithstanding that the breed is
comparatively modern, are generally acknowledged to possess great power in
impressing their likeness on all other breeds; and it is chiefly in
consequence of this power that they are so highly valued for exportation.
(14/6. Quoted by Bronn 'Geshichte der Natur' b. 2 s. 170. See Sturm 'Ueber
Racen' 1825 s. 104-107. For the niata cattle see my 'Journal of Researches'
1845 page 146.) Godine has given a curious case of a ram of a goat-like breed
of sheep from the Cape of Good Hope, which produced offspring hardly to be
distinguished from himself, when crossed with ewes of twelve other breeds. But
two of these half-bred ewes, when put to a merino ram, produced lambs closely
resembling the merino breed. Girou de Buzareingues (14/7. Lucas 'L'Heredite
Nat.' tome 2 page 112.) found that of two races of French sheep the ewes of
one, when crossed during successive generations with merino rams, yielded up
their character far sooner than the ewes of the other race. Sturm and Girou
have given analogous cases with other breeds of sheep and with cattle, the
prepotency running in these cases through the male side; but I was assured on
good authority in South America, that when niata cattle are crossed with
common cattle, though the niata breed is prepotent whether males or females
are used, yet that the prepotency is strongest through the female line. The
Manx cat is tailless and has long hind legs; Dr. Wilson crossed a male Manx
with common cats, and, out of twenty-three kittens, seventeen were destitute
of tails; but when the female Manx was crossed by common male cats all the
kittens had tails, though they were generally short and imperfect. (14/8. Mr.
Orton 'Physiology of Breeding' 1855 page 9.)
In making reciprocal crosses between pouter and fantail pigeons, the pouter-
race seemed to be prepotent through both sexes over the fantail. But this is
probably due to weak power in the fantail rather than to any unusually strong
power in the pouter, for I have observed that barbs also preponderate over
fantails. This weakness of transmission in the fantail, though the breed is an
ancient one, is said (14/9. Boitard and Corbie 'Les Pigeons' 1824 page 224.)
to be general; but I have observed one exception to the rule, namely, in a
cross between a fantail and laugher. The most curious instance known to me of
weak power in both sexes is in the trumpeter pigeon. This breed has been well
known for at least 130 years: it breeds perfectly true, as I have been assured
by those who have long kept many birds: it is characterised by a peculiar tuft
of feathers over the beak, by a crest on the head, by a singular coo quite
unlike that of any other breed, and by much-feathered feet. I have crossed
both sexes with turbits of two sub-breeds, with almond tumblers, spots, and
runts, and reared many mongrels and recrossed them; and though the crest on
the head and feathered feet were inherited (as is generally the case with most
breeds), I have never seen a vestige of the tuft over the beak or heard the
peculiar coo. Boitard and Corbie (14/10. 'Les Pigeons' pages 168, 198.) assert
that this is the invariable result of crossing trumpeters with other breeds:
Neumeister (14/11. 'Das Ganze' etc. 1837 s. 39.), however, states that in
Germany mongrels have been obtained, though very rarely, which were furnished
with the tuft and would trumpet: but a pair of these mongrels with a tuft,
which I imported, never trumpeted. Mr. Brent states (14/12. 'The Pigeon Book'
page 46.) that the crossed offspring of a trumpeter were crossed with
trumpeters for three generations, by which time the mongrels had 7/8ths of
this blood in their veins, yet the tuft over the beak did not appear. At the
fourth generation the tuft appeared, but the birds though now having 15-16ths
trumpeter's blood still did not trumpet. This case well shows the wide
difference between inheritance and prepotency; for here we have a well-
established old race which transmits its characters faithfully, but which,
when crossed with any other race, has the feeblest power of transmitting its
two chief characteristic qualities.
I will give one other instance with fowls and pigeons of weakness and strength
in the transmission of the same character to their crossed offspring. The Silk
fowl breeds true, and there is reason to believe is a very ancient race; but
when I reared a large number of mongrels from a Silk hen by a Spanish cock,
not one exhibited even a trace of the so-called silkiness. Mr. Hewitt also
asserts that in no instance are the silky feathers transmitted by this breed
when crossed with any other variety. But three birds out of many raised by Mr.
Orton from a cross between a silk cock and a bantam hen had silky feathers.
(14/13. 'Physiology of Breeding' page 22; Mr. Hewitt in 'The Poultry Book' by
Tegetmeier 1866 page 224.) So that it is certain that this breed very seldom
has the power of transmitting its peculiar plumage to its crossed progeny. On
the other hand, there is a silk sub-variety of the fantail pigeon, which has
its feathers in nearly the same state as in the Silk fowl: now we have already
seen that fantails, when crossed, possess singularly weak power in
transmitting their general qualities; but the silk sub-variety when crossed
with any other small-sized race invariably transmits its silky feathers!
(14/14. Boitard and Corbie 'Les Pigeons' 1824 page 226.)
The well-known horticulturist, Mr. Paul, informs me that he fertilised the
Black Prince hollyhock with pollen of the White Globe and the Lemonade and
Black Prince hollyhocks reciprocally; but not one seedling from these three
crosses inherited the black colour of the Black Prince. So, again, Mr. Laxton,
who has had such great experience in crossing peas, writes to me that
"whenever a cross has been effected between a white-blossomed and a purple-
blossomed pea, or between a white-seeded and a purple-spotted, brown or maple-
seeded pea, the offspring seems to lose nearly all the characteristics of the
white-flowered and white-seeded varieties; and this result follows whether
these varieties have been used as the pollen-bearing or seed-producing
parents."
The law of prepotency comes into action when species are crossed, as with
races and individuals. Gartner has unequivocally shown (14/15.
'Bastarderzeugung' s. 256, 290, etc. Naudin 'Nouvelles Archives du Museum'
tome 1 page 149 gives a striking instance of prepotency in Datura stramonium
when crossed with two other species.) that this is the case with plants. To
give one instance: when Nicotiana paniculata and vincaeflora are crossed, the
character of N. paniculata is almost completely lost in the hybrid; but if N.
quadrivalvis be crossed with N. vincaeflora, this latter species, which was
before so prepotent, now in its turn almost disappears under the power of N.
quadrivalvis. It is remarkable that the prepotency of one species over another
in transmission is quite independent, as shown by Gartner, of the greater or
less facility with which the one fertilises the other.
With animals, the jackal is prepotent over the dog, as is stated by Flourens,
who made many crosses between these animals; and this was likewise the case
with a hybrid which I once saw between a jackal and a terrier. I cannot doubt,
from the observations of Colin and others, that the ass is prepotent over the
horse; the prepotency in this instance running more strongly through the male
than through the female ass; so that the mule resembles the ass more closely
than does the hinny. (14/16. Flourens 'Longevite Humaine' page 144 on crossed
jackals. With respect to the difference between the mule and the hinny I am
aware that this has generally been attributed to the sire and dam transmitting
their characters differently; but Colin, who has given in his 'Traite Phys.
Comp.' tome 2 pages 537-539, the fullest description which I have met with of
these reciprocal hybrids, is strongly of opinion that the ass preponderates in
both crosses, but in an unequal degree. This is likewise the conclusion of
Flourens, and of Bechstein in his 'Naturgeschichte Deutschlands' b. 1 s. 294.
The tail of the hinny is much more like that of the horse than is the tail of
the mule, and this is generally accounted for by the males of both species
transmitting with greater power this part of their structure; but a compound
hybrid which I saw in the Zoological Gardens, from a mare by a hybrid ass-
zebra, closely resembled its mother in its tail.) The male pheasant, judging
from Mr. Hewitt's descriptions (14/17. Mr. Hewitt who has had such great
experience in raising these hybrids says ('Poultry Book' by Mr. Tegetmeier
1866 pages 165-167) that in all, the head was destitute of wattles, comb, and
ear-lappets; and all closely resembled the pheasant in the shape of the tail
and general contour of the body. These hybrids were raised from hens of
several breeds by a cock-pheasant; but another hybrid, described by Mr.
Hewitt, was raised from a hen-pheasant, by a silver-laced Bantam cock, and
this possessed a rudimental comb and wattles.), and from the hybrids which I
have seen, preponderates over the domestic fowl; but the latter, as far as
colour is concerned, has considerable power of transmission, for hybrids
raised from five differently coloured hens differed greatly in plumage. I
formerly examined some curious hybrids in the Zoological Gardens, between the
Penguin variety of the common duck and the Egyptian goose (Anser aegyptiacus);
and although I will not assert that the domesticated variety preponderated
over the natural species, yet it had strongly impressed its unnatural upright
figure on these hybrids.
I am aware that such cases as the foregoing have been ascribed by various
authors, not to one species, race, or individual being prepotent over the
other in impressing its character on its crossed offspring, but to such rules
as that the father influences the external characters and the mother the
internal or vital organs. But the great diversity of the rules given by
various authors almost proves their falseness. Dr. Prosper Lucas has fully
discussed this point, and has shown (14/18. 'L'Hered. Nat.' tome 2 book 2
chapter 1.) that none of the rules (and I could add others to those quoted by
him) apply to all animals. Similar rules have been announced for plants, and
have been proved by Gartner (14/19. 'Bastarderzeugung' s. 264-266. Naudin
'Nouvelles Archives du Museum' tome 1 page 148 has arrived at a similar
conclusion.) to be all erroneous. If we confine our view to the domesticated
races of a single species, or perhaps even to the species of the same genus,
some such rules may hold good; for instance, it seems that in reciprocally
crossing various breeds of fowls the male generally gives colour (14/20.
'Cottage Gardener' 1856 pages 101, 137.); but conspicuous exceptions have
passed under my own eyes. It seems that the ram usually gives its peculiar
horns and fleece to its crossed offspring, and the bull the presence or
absence of horns.
In the following chapter on Crossing I shall have occasion to show that
certain characters are rarely or never blended by crossing, but are
transmitted in an unmodified state from either parent-form; I refer to this
fact here because it is sometimes accompanied on the one side by prepotency,
which thus acquires the false appearance of unusual strength. In the same
chapter I shall show that the rate at which a species or breed absorbs and
obliterates another by repeated crosses, depends in chief part on prepotency
in transmission.]
In conclusion, some of the cases above given,--for instance, that of the
trumpeter pigeon,--prove that there is a wide difference between mere
inheritance and prepotency. This latter power seems to us, in our ignorance,
to act in most cases quite capriciously. The very same character, even though
it be an abnormal or monstrous one, such as silky feathers, may be transmitted
by different species, when crossed, either with prepotent force or singular
feebleness. It is obvious, that a purely-bred form of either sex, in all cases
in which prepotency does not run more strongly in one sex than the other, will
transmit its character with prepotent force over a mongrelised and already
variable form. (14/21. See some remarks on this head with respect to sheep by
Mr. Wilson in 'Gardener's Chronicle' 1863 page 15. Many striking instances of
this result are given by M. Malingie-Nouel 'Journ. R. Agricult. Soc.' volume
14 1853 page 220 with respect to crosses between English and French sheep. He
found that he obtained the desired influence of the English breeds by crossing
intentionally mongrelised French breeds with pure English breeds.) From
several of the above-given cases we may conclude that mere antiquity of
character does not by any means necessarily make it prepotent. In some cases
prepotency apparently depends on the same character being present and visible
in one of the two breeds which are crossed, and latent or invisible in the
other breed; and in this case it is natural that the character which is
potentially present in both breeds should be prepotent. Thus, we have reason
to believe that there is a latent tendency in all horses to be dun-coloured
and striped; and when a horse of this kind is crossed with one of any other
colour, it is said that the offspring are almost sure to be striped. Sheep
have a similar latent tendency to become dark-coloured, and we have seen with
what prepotent force a ram with a few black spots, when crossed with white
sheep of various breeds, coloured its offspring. All pigeons have a latent
tendency to become slaty-blue, with certain characteristic marks, and it is
known that, when a bird thus coloured is crossed with one of any other colour,
it is most difficult afterwards to eradicate the blue tint. A nearly parallel
case is offered by those black bantams which, as they grow old, develop a
latent tendency to acquire red feathers. But there are exceptions to the rule:
hornless breeds of cattle possess a latent capacity to reproduce horns, yet
when crossed with horned breeds they do not invariably produce offspring
bearing horns.
We meet with analogous cases with plants. Striped flowers, though they can be
propagated truly by seed, have a latent tendency to become uniformly coloured,
but when once crossed by a uniformly coloured variety, they ever afterwards
fail to produce striped seedlings. (14/22. Verlot 'Des Varietes' 1865 page
66.) Another case is in some respects more curious: plants bearing peloric
flowers have so strong a latent tendency to reproduce their normally irregular
flowers, that this often occurs by buds when a plant is transplanted into
poorer or richer soil. (14/23. Moquin-Tandon 'Teratologie' page 191.) Now I
crossed the peloric snapdragon (Antirrhinum majus), described in the last
chapter, with pollen of the common form; and the latter, reciprocally, with
peloric pollen. I thus raised two great beds of seedlings, and not one was
peloric. Naudin (14/24. 'Nouvelles Archives du Museum' tome 1 page 137.)
obtained the same result from crossing a peloric Linaria with the common form.
I carefully examined the flowers of ninety plants of the crossed Antirrhinum
in the two beds, and their structure had not been in the least affected by the
cross, except that in a few instances the minute rudiment of the fifth stamen,
which is always present, was more fully or even completely developed. It must
not be supposed that this entire obliteration of the peloric structure in the
crossed plants can be accounted for by any incapacity of transmission; for I
raised a large bed of plants from the peloric Antirrhinum, artificially
fertilised by its own pollen, and sixteen plants, which alone survived the
winter, were all as perfectly peloric as the parent-plant. Here we have a good
instance of the wide difference between the inheritance of a character and the
power of transmitting it to crossed offspring. The crossed plants, which
perfectly resembled the common snapdragon, were allowed to sow themselves, and
out of a hundred and twenty-seven seedlings, eighty-eight proved to be common
snapdragons, two were in an intermediate condition between the peloric and
normal state, and thirty-seven were perfectly peloric, having reverted to the
structure of their one grand-parent. This case seems at first sight to offer
an exception to the rule just given, namely, that a character which is present
in one form and latent in the other is generally transmitted with prepotent
force when the two forms are crossed. For in all the Scrophulariaceae, and
especially in the genera Antirrhinum and Linaria, there is, as was shown in
the last chapter, a strong latent tendency to become peloric; but there is
also, as we have seen, a still stronger tendency in all peloric plants to
reacquire their normal irregular structure. So that we have two opposed latent
tendencies in the same plants. Now, with the crossed Antirrhinums the tendency
to produce normal or irregular flowers, like those of the common Snapdragon,
prevailed in the first generation; whilst the tendency to pelorism, appearing
to gain strength by the intermission of a generation, prevailed to a large
extent in the second set of seedlings. How it is possible for a character to
gain strength by the intermission of a generation, will be considered in the
chapter on pangenesis.
On the whole, the subject of prepotency is extremely intricate,--from its
varying so much in strength, even in regard to the same character, in
different animals,--from its running either equally in both sexes, or, as
frequently is the case with animals, but not with plants, much stronger in one
sex than the other,--from the existence of secondary sexual characters,--from
the transmission of certain characters being limited, as we shall immediately
see, by sex,--from certain characters not blending together,--and, perhaps,
occasionally from the effects of a previous fertilisation on the mother. It is
therefore not surprising that no one has hitherto succeeded in drawing up
general rules on the subject of prepotency.
INHERITANCE AS LIMITED BY SEX.
New characters often appear in one sex, and are afterwards transmitted to the
same sex, either exclusively or in a much greater degree than to the other.
This subject is important, because with animals of many kinds in a state of
nature, both high and low in the scale, secondary sexual characters, not
directly connected with the organs of reproduction, are conspicuously present.
With our domesticated animals, characters of this kind often differ widely
from those distinguishing the two sexes of the parent species; and the
principle of inheritance, as limited by sex, explains how this is possible.
[Dr. P. Lucas has shown (14/25. 'L'Hered. Nat.' tome 2 pages 137-165. See also
Mr. Sedgwick's four memoirs, immediately to be referred to.) that when a
peculiarity, in no manner connected with the reproductive organs, appears in
either parent, it is often transmitted exclusively to the offspring of the
same sex, or to a much greater number of them than of the opposite sex. Thus,
in the family of Lambert, the horn-like projections on the skin were
transmitted from the father to his sons and grandsons alone; so it has been
with other cases of ichthyosis, with supernumerary digits, with a deficiency
of digits and phalanges, and in a lesser degree with various diseases,
especially with colour-blindness and the haemorrhagic diathesis, that is, an
extreme liability to profuse and uncontrollable bleeding from trifling wounds.
On the other hand, mothers have transmitted, during several generations, to
their daughters alone, supernumerary and deficient digits, colour-blindness
and other peculiarities. So that the very same peculiarity may become attached
to either sex, and be long inherited by that sex alone; but the attachment in
certain cases is much more frequent to one than the other sex. The same
peculiarities also may be promiscuously transmitted to either sex. Dr. Lucas
gives other cases, showing that the male occasionally transmits his
peculiarities to his daughters alone, and the mother to her sons alone; but
even in this case we see that inheritance is to a certain extent, though
inversely, regulated by sex. Dr. Lucas, after weighing the whole evidence,
comes to the conclusion that every peculiarity tends to be transmitted in a
greater or lesser degree to that sex in which it first appears. But a more
definite rule, as I have elsewhere shown (14/26. 'Descent of Man' 2nd edition
page 32.) generally holds good, namely, that variations which first appear in
either sex at a late period of life, when the reproductive functions are
active, tend to be developed in that sex alone; whilst variations which first
appear early in life in either sex are commonly transmitted to both sexes. I
am, however, far from supposing that this is the sole determining cause.
A few details from the many cases collected by Mr. Sedgwick (14/27. On Sexual
Limitation in Hereditary Diseases 'Brit. and For. Med.-Chirurg. Review' April
1861 page 477; July page 198; April 1863 page 445; and July page 159. Also in
1867 'On the influence of Age in Hereditary Disease.'), may be here given.
Colour-blindness, from some unknown cause, shows itself much oftener in males
than in females; in upwards of two hundred cases collected by Mr. Sedgwick,
nine-tenths related to men; but it is eminently liable to be transmitted
through women. In the case given by Dr. Earle, members of eight related
families were affected during five generations: these families consisted of
sixty-one individuals, namely, of thirty-two males, of whom nine-sixteenths
were incapable of distinguishing colour, and of twenty-nine females, of whom
only one-fifteenth were thus affected. Although colour-blindness thus
generally clings to the male sex, nevertheless, in one instance in which it
first appeared in a female, it was transmitted during five generations to
thirteen individuals, all of whom were females. The haemorrhagic diathesis,
often accompanied by rheumatism, has been known to affect the males alone
during five generations, being transmitted, however, through the females. It
is said that deficient phalanges in the fingers have been inherited by the
females alone during ten generations. In another case, a man thus deficient in
both hands and feet, transmitted the peculiarity to his two sons and one
daughter; but in the third generation,--out of nineteen grandchildren, twelve
sons had the family defect, whilst the seven daughters were free. In ordinary
cases of sexual limitation, the sons or daughters inherit the peculiarity,
whatever it may be, from their father or mother, and transmit it to their
children of the same sex; but generally with the haemorrhagic diathesis, and
often with colour-blindness, and in some other cases, the sons never inherit
the peculiarity directly from their fathers, but the daughters alone transmit
the latent tendency, so that the sons of the daughters alone exhibit it. Thus
the father, grandson, and great-great-grandson will exhibit a peculiarity,--
the grandmother, daughter, and great-grand-daughter having transmitted it in a
latent state. Hence we have, as Mr. Sedgwick remarks, a double kind of atavism
or reversion; each grandson apparently receiving and developing the
peculiarity from his grandfather, and each daughter apparently receiving the
latent tendency from her grandmother.
From the various facts recorded by Dr. Prosper Lucas, Mr. Sedgwick, and
others, there can be no doubt that peculiarities first appearing in either
sex, though not in any way necessarily or invariably connected with that sex,
strongly tend to be inherited by the offspring of the same sex, but are often
transmitted in a latent state through the opposite sex.
Turning now to domesticated animals, we find that certain characters not
proper to the parent species are often confined to, and inherited by, one sex
alone; but we do not know the history of the first appearance of such
characters. In the chapter on Sheep, we have seen that the males of certain
races differ greatly from the females in the shape of their horns, these being
absent in the ewes of some breeds; they differ also in the development of fat
in the tail and in the outline of the forehead. These differences, judging
from the character of the allied wild species, cannot be accounted for by
supposing that they have been derived from distinct parent forms. There is,
also, a great difference between the horns of the two sexes in one Indian
breed of goats. The bull zebu is said to have a larger hump than the cow. In
the Scotch deer-hound the two sexes differ in size more than in any other
variety of the dog (14/28. W. Scrope 'Art of Deer Stalking' page 354.) and,
judging from analogy, more than in the aboriginal parent-species. The peculiar
colour called tortoise-shell is very rarely seen in a male cat; the males of
this variety being of a rusty tint.
In various breeds of the fowl the males and females often differ greatly; and
these differences are far from being the same with those which distinguish the
two sexes of the parent-species, the Gallus bankiva; and consequently have
originated under domestication. In certain sub-varieties of the Game race we
have the unusual case of the hens differing from each other more than the
cocks. In an Indian breed of a white colour shaded with black, the hens
invariably have black skins, and their bones are covered by a black
periosteum, whilst the cocks are never or most rarely thus characterised.
Pigeons offer a more interesting case; for throughout the whole great family
the two sexes do not often differ much; and the males and females of the
parent-form, the C. livia, are undistinguishable: yet we have seen that with
pouters the male has the characteristic quality of pouting more strongly
developed than the female; and in certain sub-varieties the males alone are
spotted or striated with black, or otherwise differ in colour. When male and
female English carrier-pigeons are exhibited in separate pens, the difference
in the development of the wattle over the beak and round the eyes is
conspicuous. So that here we have instances of the appearance of secondary
sexual characters in the domesticated races of a species in which such
differences are naturally quite absent.]
On the other hand, secondary sexual characters which belong to the species in
a state of nature are sometimes quite lost, or greatly diminished, under
domestication. We see this in the small size of the tusks in our improved
breeds of the pig, in comparison with those of the wild boar. There are sub-
breeds of fowls, in which the males have lost the fine-flowing tail-feathers
and hackles; and others in which there is no difference in colour between the
two sexes. In some cases the barred plumage, which in gallinaceous birds is
commonly the attribute of the hen, has been transferred to the cock, as in the
cuckoo sub-breeds. In other cases masculine characters have been partly
transferred to the female, as with the splendid plumage of the golden-spangled
Hamburgh hen, the enlarged comb of the Spanish hen, the pugnacious disposition
of the Game hen, and as in the well-developed spurs which occasionally appear
in the hens of various breeds. In Polish fowls both sexes are ornamented with
a topknot, that of the male being formed of hackle-like feathers, and this is
a new male character in the genus Gallus. On the whole, as far as I can judge,
new characters are more apt to appear in the males of our domesticated animals
than in the females (14/29. I have given in my 'Descent of Man' 2nd edition
page 223 sufficient evidence that male animals are usually more variable than
the females.), and afterwards to be inherited exclusively or more strongly by
the males. Finally, in accordance with the principle of inheritance as limited
by sex, the preservation and augmentation of secondary sexual characters in
natural species offers no especial difficulty, as this would follow through
that form of selection which I have called sexual selection.
INHERITANCE AT CORRESPONDING PERIODS OF LIFE.
This is an important subject. Since the publication of my 'Origin of Species'
I have seen no reason to doubt the truth of the explanation there given of one
of the most remarkable facts in biology, namely, the difference between the
embryo and the adult animal. The explanation is, that variations do not
necessarily or generally occur at a very early period of embryonic growth, and
that such variations are inherited at a corresponding age. As a consequence of
this the embryo, even after the parent-form has undergone great modification,
is left only slightly modified; and the embryos of widely-different animals
which are descended from a common progenitor remain in many important respects
like one another and probably like their common progenitor. We can thus
understand why embryology throws a flood of light on the natural system of
classification, as this ought to be as far as possible genealogical. When the
embryo leads an independent life, that is, becomes a larva, it has to be
adapted to the surrounding conditions in its structure and instincts,
independently of those of its parents; and the principle of inheritance at
corresponding periods of life renders this possible.
This principle is, indeed, in one way so obvious that it escapes attention. We
possess a number of races of animals and plants, which, when compared with one
another and with their parent-forms, present conspicuous differences, both in
their immature and mature states. Look at the seeds of the several kinds of
peas, beans, maize, which can be propagated truly, and see how they differ in
size, colour, and shape, whilst the full-grown plants differ but little.
Cabbages, on the other hand, differ greatly in foliage and manner of growth,
but hardly at all in their seeds; and generally it will be found that the
differences between cultivated plants at different periods of growth are not
necessarily closely connected together, for plants may differ much in their
seeds and little when full-grown, and conversely may yield seeds hardly
distinguishable, yet differ much when full-grown. In the several breeds of
poultry, descended from a single species, differences in the eggs and chickens
whilst covered with down, in the plumage at the first and subsequent moults,
as well as in the comb and wattles, are all inherited. With man peculiarities
in the milk and second teeth (of which I have received the details) are
inheritable, and longevity is often transmitted. So again with our improved
breeds of cattle and sheep, early maturity, including the early development of
the teeth, and with certain breeds of fowl the early appearance of secondary
sexual characters, all come under the same head of inheritance at
corresponding periods.
Numerous analogous facts could be given. The silk-moth, perhaps, offers the
best instance; for in the breeds which transmit their characters truly, the
eggs differ in size, colour, and shape: the caterpillars differ, in moulting
three or four times, in colour, even in having a dark-coloured mark like an
eyebrow, and in the loss of certain instincts;--the cocoons differ in size,
shape, and in the colour and quality of the silk; these several differences
being followed by slight or barely distinguishable differences in the mature
moth.
But it may be said that, if in the above cases a new peculiarity is inherited,
it must be at the corresponding stage of development; for an egg or seed can
resemble only an egg or seed, and the horn in a full-grown ox can resemble
only a horn. The following cases show inheritance at corresponding periods
more plainly, because they refer to peculiarities which might have supervened,
as far as we can see, earlier or later in life, yet are inherited at the same
period at which they first appeared.
[In the Lambert family the porcupine-like excrescences appeared in the father
and sons at the same age, namely, about nine weeks after birth. (14/30.
Prichard 'Phys. Hist. of Mankind' 1851 volume 1 page 349.) In the
extraordinary hairy family described by Mr. Crawfurd (14/31. 'Embassy to the
Court of Ava' volume 1 page 320. The third generation is described by Capt.
Yule in his 'Narrative of the Mission to the Court of Ava' 1855 page 94.),
children were produced during three generations with hairy ears; in the father
the hair began to grow over his body at six years old; in his daughter
somewhat earlier, namely, at one year; and in both generations the milk teeth
appeared late in life, the permanent teeth being afterwards singularly
deficient. Greyness of hair at an unusually early age has been transmitted in
some families. These cases border on diseases inherited at corresponding
periods of life, to which I shall immediately refer.
It is a well-known peculiarity with almond-tumbler pigeons, that the full
beauty and peculiar character of the plumage does not appear until the bird
has moulted two or three times. Neumeister describes and figures a brace of
pigeons in which the whole body is white except the breast, neck, and head;
but in their first plumage all the white feathers have coloured edges. Another
breed is more remarkable: its first plumage is black, with rusty-red wing-bars
and a crescent-shaped mark on the breast; these marks then become white, and
remain so during three or four moults; but after this period the white spreads
over the body, and the bird loses its beauty. (14/32. 'Das Ganze der
Taubenzucht' 1837 s. 24 tab. 4 figure 2 s. 21 tab. 1 figure 4.) Prize canary-
birds have their wings and tail black: "this colour, however, is only retained
until the first moult, so that they must be exhibited ere the change takes
place. Once moulted, the peculiarity has ceased. Of course all the birds
emanating from this stock have black wings and tails the first year." (14/33.
Kidd 'Treatise on the Canary' page 18.) A curious and somewhat analogous
account has been given (14/34. Charlesworth 'Mag. of Nat. Hist.' volume 1 1837
page 167.) of a family of wild pied rooks which were first observed in 1798,
near Chalfont, and which every year from that date up to the period of the
published notice, viz., 1837 "have several of their brood particoloured, black
and white. This variegation of the plumage, however, disappears with the first
moult; but among the next young families there are always a few pied ones."
These changes of plumage, which are inherited at various corresponding periods
of life in the pigeon, canary-bird, and rook, are remarkable, because the
parent-species passes through no such change.
Inherited diseases afford evidence in some respects of less value than the
foregoing cases, because diseases are not necessarily connected with any
change in structure; but in other respects of more value, because the periods
have been more carefully observed. Certain diseases are communicated to the
child apparently by a process like inoculation, and the child is from the
first affected; such cases may be here passed over. Large classes of diseases
usually appear at certain ages, such as St. Vitus's dance in youth,
consumption in early mid-life, gout later, and apoplexy still later; and these
are naturally inherited at the same period. But even in diseases of this
class, instances have been recorded, as with St. Vitus's dance, showing that
an unusually early or late tendency to the disease is inheritable. (14/35. Dr.
Prosper Lucas 'Hered. Nat.' tome 2 page 713.) In most cases the appearance of
any inherited disease is largely determined by certain critical periods in
each person's life, as well as by unfavourable conditions. There are many
other diseases, which are not attached to any particular period, but which
certainly tend to appear in the child at about the same age at which the
parent was first attacked. An array of high authorities, ancient and modern,
could be given in support of this proposition. The illustrious Hunter believed
in it; and Piorry (14/36. 'L'Hered. dans les Maladies' 1840 page 135. For
Hunter see Harlan 'Med. Researches' page 530.) cautions the physician to look
closely to the child at the period when any grave inheritable disease attacked
the parent. Dr. Prosper Lucas (14/37. 'L'Hered. Nat.' tome 2 page 850.), after
collecting facts from every source, asserts that affections of all kinds,
though not related to any particular period of life, tend to reappear in the
offspring at whatever period of life they first appeared in the progenitor.
As the subject is important, it may be well to give a few instances, simply as
illustrations, not as proof; for proof, recourse must be had to the
authorities above quoted. Some of the following cases have been selected for
the sake of showing that, when a slight departure from the rule occurs, the
child is affected somewhat earlier in life than the parent. In the family of
Le Compte blindness was inherited through three generations, and no less than
twenty-seven children and grandchildren were all affected at about the same
age; their blindness in general began to advance about the fifteenth or
sixteenth year, and ended in total deprivation of sight at the age of about
twenty-two. (14/38. Sedgwick 'Brit. and For. Med.-Chirurg. Review' April 1861
page 485. In some accounts the number of children and grandchildren is given
as 37; but this seems to be an error judging from the paper first published in
the 'Baltimore Med. and Phys. Reg.' 1809 of which Mr. Sedgwick has been so
kind as to send me a copy.) In another case a father and his four children all
became blind at twenty-one years old; in another, a grandmother grew blind at
thirty-five, her daughter at nineteen, and three grandchildren at the ages of
thirteen and eleven. (14/39. Prosper Lucas 'Hered. Nat.' tome 1 page 400.) So
with deafness, two brothers, their father and paternal grandfather, all became
deaf at the age of forty. (14/40. Sedgwick ibid July 1861 page 202.)
Esquirol gives several striking instances of insanity coming on at the same
age, as that of a grandfather, father, and son, who all committed suicide near
their fiftieth year. Many other cases could be given, as of a whole family who
became insane at the age of forty. (14/41. Piorry page 109; Prosper Lucas tome
2 page 759.) Other cerebral affections sometimes follow the same rule,--for
instance, epilepsy and apoplexy. A woman died of the latter disease when
sixty-three years old; one of her daughters at forty-three, and the other at
sixty-seven: the latter had twelve children, who all died from tubercular
meningitis. (14/42. Prosper Lucas tome 2 page 748.) I mention this latter case
because it illustrates a frequent occurrence, namely, a change in the precise
nature of an inherited disease, though still affecting the same organ.
Asthma has attacked several members of the same family when forty years old,
and other families during infancy. The most different diseases, such as angina
pectoris, stone in the bladder, and various affections of the skin, have
appeared in successive generations at nearly the same age. The little finger
of a man began from some unknown cause to grow inwards, and the same finger in
his two sons began at the same age to bend inwards in a similar manner.
Strange and inexplicable neuralgic affections have caused parents and children
to suffer agonies at about the same period of life. (14/43. Prosper Lucas tome
3 pages 678, 700, 702; Sedgwick ibid April 1863 page 449 and July 1863 page
162. Dr. J. Steinan 'Essay on Hereditary Disease' 1843 pages 27, 34.)
I will give only two other cases, which are interesting as illustrating the
disappearance as well as the appearance of disease at the same age. Two
brothers, their father, their paternal uncles, seven cousins, and their
paternal grandfather, were all similarly affected by a skin-disease, called
pityriasis versicolor; "the disease, strictly limited to the males of the
family (though transmitted through the females), usually appeared at puberty,
and disappeared at about the age of forty or forty-five years." The second
case is that of four brothers, who when about twelve years old suffered almost
every week from severe headaches, which were relieved only by a recumbent
position in a dark room. Their father, paternal uncles, paternal grandfather,
and granduncles all suffered in the same way from headaches, which ceased at
the age of fifty-four or fifty-five in all those who lived so long. None of
the females of the family were affected. (14/44. These cases are given by Mr.
Sedgwick on the authority of Dr. H. Stewart in 'Med.-Chirurg. Review' April
1863 pages 449, 477.)]
It is impossible to read the foregoing accounts, and the many others which
have been recorded, of diseases coming on during three or even more
generations in several members of the same family at the same age, especially
in the case of rare affections in which the coincidence cannot be attributed
to chance, and to doubt that there is a strong tendency to inheritance in
disease at corresponding periods of life. When the rule fails, the disease is
apt to come on earlier in the child than in the parent; the exceptions in the
other direction being very much rarer. Dr. Lucas (14/45. 'Hered. Nat.' tome 2
page 852.) alludes to several cases of inherited diseases coming on at an
earlier period. I have already given one striking instance with blindness
during three generations; and Mr. Bowman remarks that this frequently occurs
with cataract. With cancer there seems to be a peculiar liability to earlier
inheritance: Sir J. Paget, who has particularly attended to this subject, and
tabulated a large number of cases, informs me that he believes that in nine
cases out of ten the later generation suffers from the disease at an earlier
period than the previous generation. He adds, "In the instances in which the
opposite relation holds, and the members of later generations have cancer at a
later age than their predecessors, I think it will be found that the non-
cancerous parents have lived to extreme old ages." So that the longevity of a
non-affected parent seems to have the power of influencing the fatal period in
the offspring; and we thus apparently get another element of complexity in
inheritance.
The facts, showing that with certain diseases the period of inheritance
occasionally or even frequently advances, are important with respect to the
general descent-theory, for they render it probable that the same thing would
occur with ordinary modifications of structure. The final result of a long
series of such advances would be the gradual obliteration of characters proper
to the embryo and larva, which would thus come to resemble more and more
closely the mature parent-form. But any structure which was of service to the
embryo or larva would be preserved by the destruction at this stage of growth
of each individual which manifested any tendency to lose its proper character
at too early an age.
Finally, from the numerous races of cultivated plants and domestic animals, in
which the seeds or eggs, the young or old, differ from one another and from
those of the parent-species;--from the cases in which new characters have
appeared at a particular period, and afterwards been inherited at the same
period;--and from what we know with respect to disease, we must believe in the
truth of the great principle of inheritance at corresponding periods of life.
SUMMARY OF THE THREE PRECEDING CHAPTERS.
Strong as is the force of inheritance, it allows the incessant appearance of
new characters. These, whether beneficial or injurious,--of the most trifling
importance, such as a shade of colour in a flower, a coloured lock of hair, or
a mere gesture,--or of the highest importance, as when affecting the brain, or
an organ so perfect and complex as the eye,--or of so grave a nature as to
deserve to be called a monstrosity,--or so peculiar as not to occur normally
in any member of the same natural class,--are often inherited by man, by the
lower animals, and plants. In numberless cases it suffices for the inheritance
of a peculiarity that one parent alone should be thus characterised.
Inequalities in the two sides of the body, though opposed to the law of
symmetry, may be transmitted. There is ample evidence that the effects of
mutilations and of accidents, especially or perhaps exclusively when followed
by disease, are occasionally inherited. There can be no doubt that the evil
effects of the long-continued exposure of the parent to injurious conditions
are sometimes transmitted to the offspring. So it is, as we shall see in a
future chapter, with the effects of the use and disuse of parts, and of mental
habits. Periodical habits are likewise transmitted, but generally, as it would
appear, with little force.
Hence we are led to look at inheritance as the rule, and non-inheritance as
the anomaly. But this power often appears to us in our ignorance to act
capriciously, transmitting a character with inexplicable strength or
feebleness. The very same peculiarity, as the weeping habit of trees, silky
feathers, etc., may be inherited either firmly or not at all by different
members of the same group, and even by different individuals of the same
species, though treated in the same manner. In this latter case we see that
the power of transmission is a quality which is merely individual in its
attachment. As with single characters, so it is with the several concurrent
slight differences which distinguish sub-varieties or races; for of these,
some can be propagated almost as truly as species, whilst others cannot be
relied on. The same rule holds good with plants, when propagated by bulbs,
offsets, etc., which in one sense still form parts of the same individual, for
some varieties retain or inherit through successive bud-generations their
character far more truly than others.
Some characters not proper to the parent-species have certainly been inherited
from an extremely remote epoch, and may therefore be considered as firmly
fixed. But it is doubtful whether length of inheritance in itself gives
fixedness of character; though the chances are obviously in favour of any
character which has long been transmitted true or unaltered still being
transmitted true as long as the conditions of life remain the same. We know
that many species, after having retained the same character for countless
ages, whilst living under their natural conditions, when domesticated have
varied in the most diversified manner, that is, have failed to transmit their
original form; so that no character appears to be absolutely fixed. We can
sometimes account for the failure of inheritance by the conditions of life
being opposed to the development of certain characters; and still oftener, as
with plants cultivated by grafts and buds, by the conditions causing new and
slight modifications incessantly to appear. In this latter case it is not that
inheritance wholly fails, but that new characters are continually superadded.
In some few cases, in which both parents are similarly characterised,
inheritance seems to gain so much force by the combined action of the two
parents, that it counteracts its own power, and a new modification is the
result.
In many cases the failure of the parents to transmit their likeness is due to
the breed having been at some former period crossed; and the child takes after
his grandparent or more remote ancestor of foreign blood. In other cases, in
which the breed has not been crossed, but some ancient character has been lost
through variation, it occasionally reappears through reversion, so that the
parents apparently fail to transmit their own likeness. In all cases, however,
we may safely conclude that the child inherits all its characters from its
parents, in whom certain characters are latent, like the secondary sexual
characters of one sex in the other. When, after a long succession of bud-
generations, a flower or fruit becomes separated into distinct segments,
having the colours or other attributes of both parent-forms, we cannot doubt
that these characters were latent in the earlier buds, though they could not
then be detected, or could be detected only in an intimately commingled state.
So it is with animals of crossed parentage, which with advancing years
occasionally exhibit characters derived from one of their two parents, of
which not a trace could at first be perceived. Certain monstrosities, which
resemble what naturalists call the typical form of the group in question,
apparently come under the same law of reversion. It is assuredly an
astonishing fact that the male and female sexual elements, that buds, and even
full-grown animals, should retain characters, during several generations in
the case of crossed breeds, and during thousands of generations in the case of
pure breeds, written as it were in invisible ink, yet ready at any time to be
evolved under certain conditions.
What these conditions precisely are, we do not know. But any cause which
disturbs the organisation or constitution seems to be sufficient. A cross
certainly gives a strong tendency to the reappearance of long-lost characters,
both corporeal and mental. In the case of plants, this tendency is much
stronger with those species which have been crossed after long cultivation and
which therefore have had their constitutions disturbed by this cause as well
as by crossing, than with species which have always lived under their natural
conditions and have then been crossed. A return, also, of domesticated animals
and cultivated plants to a wild state favours reversion; but the tendency
under these circumstances has been much exaggerated.
When individuals of the same family which differ somewhat, and when races or
species are crossed, the one is often prepotent over the other in transmitting
its character. A race may possess a strong power of inheritance, and yet when
crossed, as we have seen with trumpeter-pigeons, yield to the prepotency of
every other race. Prepotency of transmission may be equal in the two sexes of
the same species, but often runs more strongly in one sex. It plays an
important part in determining the rate at which one race can be modified or
wholly absorbed by repeated crosses with another. We can seldom tell what
makes one race or species prepotent over another; but it sometimes depends on
the same character being present and visible in one parent, and latent or
potentially present in the other.
Characters may first appear in either sex, but oftener in the male than in the
female, and afterwards be transmitted to the offspring of the same sex. In
this case we may feel confident that the peculiarity in question is really
present though latent in the opposite sex! hence the father may transmit
through his daughter any character to his grandson; and the mother conversely
to her granddaughter. We thus learn, and the fact is an important one, that
transmission and development are distinct powers. Occasionally these two
powers seem to be antagonistic, or incapable of combination in the same
individual; for several cases have been recorded in which the son has not
directly inherited a character from his father, or directly transmitted it to
his son, but has received it by transmission through his non-affected mother,
and transmitted it through his non-affected daughter. Owing to inheritance
being limited by sex, we see how secondary sexual characters may have arisen
under nature; their preservation and accumulation being dependent on their
service to either sex.
At whatever period of life a new character first appears, it generally remains
latent in the offspring until a corresponding age is attained, and then is
developed. When this rule fails, the child generally exhibits the character at
an earlier period than the parent. On this principle of inheritance at
corresponding periods, we can understand how it is that most animals display
from the germ to maturity such a marvellous succession of characters.
Finally, though much remains obscure with respect to Inheritance, we may look
at the following laws as fairly well established.
FIRSTLY, a tendency in every character, new and old, to be transmitted by
seminal and bud generation, though often counteracted by various known and
unknown causes.
SECONDLY, reversion or atavism, which depends on transmission and development
being distinct powers: it acts in various degrees and manners through both
seminal and bud generation.
THIRDLY, prepotency of transmission, which may be confined to one sex, or be
common to both sexes.
FOURTHLY, transmission, as limited by sex, generally to the same sex in which
the inherited character first appeared; and this in many, probably most cases,
depends on the new character having first appeared at a rather late period of
life.
FIFTHLY, inheritance at corresponding periods of life, with some tendency to
the earlier development of the inherited character.
In these laws of Inheritance, as displayed under domestication, we see an
ample provision for the production, through variability and natural selection,
of new specific forms.